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ILLINOIS  BIOLOGICAL 
MONOGRAPHS 


PUBLISHED  QUARTERLY 

UNDER  THE  AUSPICES  OF  THE  GRADUATE  SCHOOL 

BY  THE  UNIVERSITY  OF  ILLINOIS 


VOLUME  I 


Urbana,  Illinois 
1914-15 


Editorial  Committee 


Stephen  Alfred  Forbes  William  Trelease 

Henry  Baldwin  Ward 


/ 


7 


TABLE  OF  CONTENTS 

Volume  I 


NUMBERS  PAGES 

I  and  2    A  Revision  of  the  Cestode  Family  Proteocephalidae.    By  George 

Roger  La  Rue.    With  i6  plates 1-350 

(Distributed  November  30,  1914) 

3        Studies   on    the    Cestode   Family   Anoplocephalidae.     By    Herman 

Douthitt.    With  6  plates 351-446 

(Distributed  March  8,  1915) 


4        Some   North  American  Larval   Trematodes.     By  William   Walter 

Cort.    With  8  plates 447-532 

(Distributed  June  28,  1915) 


732170 


UNIVERSITY      OF      ILLINOIS      BULLETIN 


VOL..    XI  i 

[Euterc.i 


Ilie    .\i-t    ■■!     .Ailt;'.!"!    J4.     171J.I 


ILLINOIS  BIOLOGICAL 
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July-October,  1914  Nos.  1  and  2 


A  REVISION  OI^  THE  CESTODE  FAMILY 
PROTEOCEPHALIDiE 


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ILLINOIS    BIOLOGICAL 
MONOGRAPHS 


Vol.  I  July-October,  1914  Nos.  1  and  2 


Editorial  Committee 


Stephen  Alfred  Forbes  William  Trelease 

Henry  Baldwin  Ward 


Published  by  the 

University  of  Illinois 

Urbana 


Copyright,  1914 
By  the  University  of  Illinois 


A   REVISION  OF 
THE    CESTODE    FAMILY 

PROTEOCEPHALID^ 


BY 


GEORGE  ROGER  LA  RUE 


Contributions  from  the 

Zoological   Laboratory   of  the   University   of   Illinois   under  the  direction   of 

Henry  B.  Ward,  No.  33 


THESIS 

Submitted  in  Partial  Fulfilment  of  the  Requirements 

for  the  Degree  of  Doctor  of  Philosophy 

in  Zoology  in  the  Graduate  School 

of  the  University  of  Illinois 

1911 


TABLE  OF  CONTENTS 

PAGE 

Introduction  _ 7 

Historical  — 9 

Synonymy    _ lo 

Deiinitions  of  Families  and  Genera  Considered ii 

Methods   of   Technique 14 

Anatomy  and  Histology  of  Proteocephalids 16 

Characters  of  Diagnostic  Value 33 

Key  to  the  Better  Known  Genera  and  Species  of  Proteocephalidae 34 

Description   of   Proteocephalid  Species yj 

Proteocephalus  filicoUis  ( Rudolphi ) 38 

P.  esocis   (G.   Schneider) 48 

P.  agonis  (Barbieri) 51 

P.  exiguus  La  Rue 54 

P.  pusillus  Ward 58 

P.  pinguis  La  Rue 65 

P.  fallax  La  Rue. 70 

P.  neglectus  La  Rue „ 78 

P.  dubius  La  Rue 81 

P.  cernuae  (Gmelin)  La  Rue. 86 

P.  percae  (  Miiller ) 93 

Table  Showing  Comparative  Data  on  P.  ocellatus  and  P.  percae....  106 

P.  longicollis  ( Zeder ) 109 

P.  torulosus  (  Batsch  ) 1 18 

P.  macrocephalus  (Creplin) 129 

P.  ambloplitis  ( Leidy ) „ I43 

P.  perplexus  La  Rue iSi 

P.  singularis  La  Rue ~ 156 

P.  sulcatus   ( Klaptocz) 162 

P.  pentastoma   ( Klaptocz) 165 

P.  fossatus  (Riggenbach)  167 

P.  skorikowi  {von  Linstow) 169 

P.  sagittus   (Grimm)   171 

P.  salvelini  (Linton) 174 

P.  Cyclops  (von  Linstow)   sp.  inq I75 

P.  hemisphericus  (Molin)  sp.  inq 175 

P.  macro  phallus  (Diesing)  sp.  inq I77 

P.  nematosoma   (Leidy)    sp.  inq 177 

P.  salmonis-umblae  (Monticelli)  sp.  inq I79 

P.  osculatus  (Goeze)  sp.  inq 180 

Taenia  siwplicissima  Leidy  186 

Taenia  belones  Mullcr 187 


6  iiUNOlS  BIOLOGICAL  MONOGRAPHS  [6 

PAGE 

Taenia  pollachii  Rathke 187 

Choanoscolex  abscisus  (Riggenbach)  La  Rue 188 

G>mparative  Table  of  Selected  Characters  of  Proteocephalus  Species 192 

^    Description  of  Proteocephalid  Species  from  Amphibia  and  Reptiles : 

Ophiotaenia  perspicua  La  Rue 204 

O.  lactea  (Leidy)  sp.  inq 208 

O.  filaroides  La  Rue „ 209 

O.  lonnbergii  (Fuhrmann)  214 

O.  nattereri    (Parona)    218 

O.  racemosa    (Rudolphi) 220 

O.  pigmentata    (von  Linstow)    226 

O.  trimeresuri    (Parona) „ 227 

O.  coitnettei    (Barrois)    „ _ 231 

O.  punica   (Cholodkovski)   „ 237 

O,  marenselleri    (  Barrois) „ 240 

O.  grandis  La  Rue. 242 

Crepidobothriutn   gerrardii    ( Baird) 246 

Comparative  Tables   of   Selected    Characters  of   Species  of    Ophiotaenia  and 

Crepidobothriutn  260 

Oochoristica  cryptobothrium  (von  Linstow)  La  Rue 268 

^  Description  of  Species  of  Monticellia 

Monticellia  coryphicephala  (Monticelli)  La  Rue 268 

M.  macrocotylea  (Monticelli) _ 275 

M.  diesingii  (Monticelli) 276 

M.  malopteruri  (Fritsch)  La  Rue 277 

Distribution    of    Proteocephalidae _ _ 281 

Distribution  of   Proteocephalids  of  Amphibia  and  Reptilia 281 

Hosts  and  Localities  Reported   for   Proteocephalids  of  Amphibia 

and  Reptilia  _ 283 

Families    of    Amphibia    and    Reptilia    Harboring    Proteocephalid 

Species  _ _ 284 

Hosts  and  Localities  of  Proteocephalid  Species  Infesting  Fish  and 

the  Species  of  Fish  Infested „ 285 

Distribution  of  the  Proteocephalids  of  Fish  According  to  Continents  288 

Fish  Harboring  Two  or  More  Proteocephalid  Species 289 

Proteocephalids  Occurring  in  Two  or  More  Species  of  Fish 290 

Families  of  Fish  from  which  Proteocephalids  are  known _  305 

Life  History  of  the    Proteocephalidae ~ ™  306 

Origin  of  the  Proteocephalidae 307 

Bibliography    _ 310 

Explanation  of  Plates  « _ 3^9 


7]  PROTEOCEPHALIDAE  —  LA  RUE 


INTRODUCTION 

Several  years  ago  while  studying  under  Professor  Henry  B.  Ward 
at  the  University  of  Nebraska  the  writer  began  an  investigation  of  a 
<;estode  parasitic  in  Ambly stoma  tigrinum  (Green).  That  investigation 
resulted  in  a  paper  (La  Rue  1909)  in  which  a  number  of  points  in  the 
anatomy  of  Proteocephalids  were  cleared  up  and  certain  problems  were 
outlined  for  investigation  at  an  early  date.  In  the  meantime  the  writer 
became  interested  in  the  large  number  of  Proteocephalids  which  Pro- 
fessor Ward  had  secured  by  work  in  the  field  and  by  exchange.  The 
writer  had  also  made  some  collections.  A  preliminary  study  of  the 
available  material  by  the  writer  was  convincing  to  Professor  Ward  that 
a  more  complete  and  comparative  study  of  the  group  was  desirable  and 
that  the  results  of  such  a  study  would  prove  of  value  to  helminthologists, 
not  only  of  America  but  also  in  Europe.  Such  a  study  seemed  more 
desirable  since  but  two  comparative  studies  of  the  group  had  been  made 
in  more  than  fifteen  years,  the  first  by  Riggenbach  (1896)  mostly  on  the 
species  infesting  fish,  and  the  second  by  Schwarz  (1908)  wholly  on  the 
species  infesting  snakes  and  reptiles.  The  fact  that  very  little  had 
been  done  on  the  genus  by  American  investigators  furnished  an  added 
incentive  to  undertake  the  work. 

At  Professor  Ward's  suggestion  and  under  his  direction  the  writer 
"undertook  an  investigation  of  the  genus  which  would  be  comprehensive 
in  scope  and  at  the  same  time  comparative.  For  this  purpose  Professor 
Ward  augmented  the  material  already  at  hand  by  securing  the  loan  of 
alcoholics  and  prepared  slides  from  several  European  and  American 
investigators  and  from  the  collections  of  the  Smithsonian  Institution 
and  the  Bureau  of  Animal  Industry  at  Washington.  Unfortunately 
specimens  of  certain  material  could  not  be  secured  either  because  it 
had  been  lost  or  because  it  could  not  be  removed  from  the  collections. 

The  lack  of  some  of  this  material  has  caused  the  writer  to  change 
the  plan  of  the  paper  somewhat.  Instead  of  making  a  fresh  study  of 
each  species  it  has  been  necessary  in  describing  certain  species  to  depend 
on  the  data  secured  from  the  literature.  These  data  have  been  recom- 
liined  and  quotations  have  been  made,  sometimes  quite  extended,  while 


8  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [S 

original  drawings  which  were  found  useful  in  presenting  the  characters 
and  structures  of  the  species  have  been  copied  and  used  in  the  new 
account.  Possibly  it  may  seem  that  a  consideration  of  these  species 
might  have  been  omitted  without  apparently  having  great  effect  on  the 
value  of  the  work.  The  writer  found  however  that  serious  errors  had 
been  made  in  determination  and  upon  these  mistaken  determinations 
identification  of  other  species  had  been  made  dependent.  The  only 
way  of  escape  from  the  tangle  was  to  consider  every  species  in  the  family. 

In  order  that  the  work  might  be  of  more  value  to  many  of  the 
American  investigators  to  whom  large  numbers  of  the  older  works  of  the 
European  writers  are  unavailable  and  for  the  purpose  of  comparison 
large  extracts  from  these  older  writings  have  been  quoted  verbatim. 
For  these  reasons  also  the  number  of  drawings  copied  is  larger  than  at 
first  glimpse  seems  necessary.  The  writer  has  been  compelled  however 
to  omit  because  of  lack  of  space  many  extracts  and  many  drawings 
which  would  add  to  the  value  of  the  work.  In  working  over  the  large 
literature  on  the  subject  an  attempt  was  made  to  secure  every  important 
paper.    Some  less  important  papers  are  known  to  have  been  omitted. 

Thanks  are  due  to  the  following  investigators  who  at  the  request 
of  Professor  "Ward  so  kindly  sent  prepared  slides  or  alcoholics  from  their 
valuable  collections  for  study  and  comparison :  Professor  Fritz 
Zschokke,  University  of  Basle,  Professor  Corrado  Parona,  University  of 
Genoa,  Professor  Max  Braun,  University  of  Konigsberg,  Professor  K.  M. 
Levander,  Helsingfors,  Finland,  Professor  Anton  Collin,  University  of 
Berlin,  Professor  A.  E.  Shipley,  Christ's  CoUege,  Cambridge,  Professor 
Fr.  Sav.  Monticelli,  University  of  Naples,  Professor  0.  Fuhrmann,  Uni- 
versity of  Neuchatel,  Dr.  C.  W.  Stiles,  Hygienic  Laboratory,  Washing- 
ton, D.  C,  Dr.  B.  H.  Bansom,  Bureau  of  Animal  Industry,  Washington, 
D.  C,  Professor  Edwin  Linton,  Washington  and  Jefferson  College, 
Professor  A.  J.  Smith,  University  of  Pennsylvania,  Professor  L.  T. 
Hankinson,  Charleston,  111.  Mr.  E.  G.  Davis  of  Lincoln,  Nebr.,  and 
Mr.  Herman  Douthitt  of  Sulphur,  Oklahoma,  have  also  very  kindly 
furnished  material  for  study.  Thanks  are  due  Mr,  S.  Fred  Prince  for 
re-drawing  many  of  the  figures  from  other  works  and  for  making  my 
pencil  sketches  into  the  finished  drawings. 

To  the  United  States  Bureau  of  Fisheries  and  to  its  corps  of  scien- 
tific workers  I  am  indebted  for  encouragement  and  assistance  in  many 
ways,  especially  in  securing  valuable  material  from  various  sources. 

To  Professor  Henry  B,  Ward  my  sincerest  thanks  are  due  for  the 
use  of  his  extensive  collections,  for  the  use  of  his  library,  and  also  for 
securing  for  my  use  many  rare  specimens  and  rare  books.  His  interest 
and  co-operation  have  given  me  the  inspiration  to  complete  the  work. 


9]  PROTEOCEPHAUDAE  —  LA  RUE 


HISTORICAL  DATA 

Rudolphi  (1808-10)  collected  together  the  results  of  the  systematic 
labors  of  helminthologists  up  to  that  time.  His  work  is  very  complete 
and  in  it  are  to  be  found  the  diagnoses,  synonymy  and  descriptions  of 
the  species  of  this  genus  that  were  known  at  that  time.  These  species 
are  included  in  the  genus  Taenia  Linnaeus  1758.  It  is  to  be  noted  that 
Rudolphi  renamed  certain  of  these  species  which  had  been  previously 
established  by  other  investigators.  Later  Rudolphi  (1819)  brought 
down  to  date  a  summary  of  the  investigations  of  preceding  helmin- 
thologists. 

Some  of  the  early  investigators,  other  than  Rudolphi,  who  reported 
species  now  included  in  the  genus  Proteocephalus,  were  Goeze,  0.  F. 
Miiller,  Pallas,  Gmelin,  Batsch,  Schrank,  Zeder,  Bloch,  and  Froelich. 
Many  of  their  descriptions  and  diagnoses  were  very  short.  Only  a  few 
were  accompanied  by  drawings.  It  is  needless  to  say  that  the  works 
of  these  early  investigators  are  almost  entirely  concerned  with  the  study 
of  external  characters.  There  were  no  more  important  works  which 
have  to  do  with  this  group  of  cestodes  until  the  time  of  Dujardin  (1845) 
and  Diesing  (1850).  These  investigators  listed  several  new  species  and 
new  host  species. 

Investigators  up  to  the  time  of  Monticelli  (1891)  with  the  excep- 
tion of  Weinland  (1858)  considered  this  group  of  fish  cestodes  to  belong 
to  the  genus  Taenia.  The  latter  author  proposed  the  name  Proteocepha- 
lus which  will  be  discussed  later  and  the  former  proposed  the  name 
Tetracotylus.  It  was  von  Linstow  (1891)  who  first  pointed  out  that 
the  fish  species  made  up  a  closely  related  group  within  the  genus  Taenia. 
He  made  the  first  careful  study  of  one  of  the  species  of  this  group. 

Monticelli  (1891)  made  a  careful  study  of  the  group  and  pointed 
out  several  misinterpretations  relating  to  the  genital  organs.  He  first 
showed  the  true  relationships  of  the  organs  in  the  interovarial 
space.  His  work  was  largely  based  on  Taenia  coryphicephala  with 
which  he  compared  other  species.  In  all  he  listed  20  species  belonging 
to  his  genus  Tetracotylus.  Of  these  species  some  were  from  snakes. 
For  a  more  complete  discussion  of  that  work  see  the  description  of 
Monticellia  (Tetracotylus).  Since  this  work  of  Monticelli,  Lonnberg 
(1894)  proposed  the  genus  Ich  thy  ©taenia  and  in  this  genus  many  species 
have  been  included.  The  range  of  hosts  includes  amphibians,  snakes, 
lizards,  and  all  the  larger  groups  of  freshwater  fish. 


10  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [10 

In  1899  Monticelli  recognized  that  Tetrabothrium  gerrardii  Baird, 
a  parasite  of  the  Boidae,  belonged  in  this  family  of  cestodes  but  that  it 
should  be  separated  from  the  species  of  Proteocephalus,  and  for  this 
species  he  proposed  the  genus  Crepidobothrium.  In  1903  von  Linstow 
proposed  the  genus  Acanthotaenia  to  include  a  spiny  headed  Proteo- 
cephalid  from  Varanus.  This  genus  was  later  emended  by  Johnston 
{1909)  who  placed  three  other  species  in  it,  all  from  Varanidae. 

It  has  been  found  necessary  to  restrict  the  genus  Proteocephalus  by 
removing  from  it  the  species  of  amphibian  and  ophidian  Proteocepha- 
lids.  These  make  a  fairly  homogeneous  group  for  which  the  writer 
proposed  the  name  Ophiotaenia  La  Rue  1911  with  Ophiotaenia  perspicua 
Xia  Rue  as  the  type  species. 

SYNONYMY 

Weinland  (1858:53)  proposed  the  genus  Proteocephalus  and  named 
as  its  type  Taenia  ambigua  Dujardin.  Taenia  filicollis  and  Taenia  dis- 
par  were  also  included  under  this  genus. 

Lonnberg  (1894:801-803)  proposed  the  generic  name  Ichthyotae- 
nia  and  listed  as  members  of  this  genus:  'Z.  filicollis  Rud.,  I.  ocellata 
Rud.,  7.  longicollis  Rud.,  I.  torulosa  Batsch,  and  /.  coryphicephala  Mon- 
ticelli. Since  Lonnberg  named  I.  filicollis  first  in  his  list  that  name  is 
to  be  considered  the  type  of  his  genus  and  has  been  so  designated  by 
Hall  (1910).  Thus  he  made  Ichthyotaenia  a  synonym  of  Proteocepha- 
lus unless  T.  filicollis  Rud.  is  genericaUy  different  from  T.  ambigua 
(Dujardin).  On  this  point  Railliet  (1899)  says:  "Comme  cette  espece 
(Taenia  ambigua  Duj.)  re^tre  nettement  dans  le  genre  Ichthyotaenia  il 
est  evident  que  le  premier  nom  (Proteocephalus)  doit  etre  repris." 
The  validity  of  the  names  Proteocephalus  Weinland  or  Ichthyotaenia 
Lonnberg  then  depends  on  the  species  T.  ambigua  Dujardin  and  T. 
filicollis  Rud.  The  first  named  species  was  fairly  well  described  and 
measurements  of  diagnostic  value  were  given  by  Dujardin  (1845).  This 
species  has  since  been  well  described  and  figured  by  G.  Schneider  and 
later  by  the  writer  using  Schneider's  material  which  agrees  almost  per- 
fectly with  Dujardin 's  description  of  T.  ambigua. 

Taenia  filicollis  Rud.  is  not  a  synonym  of  Taenia  ocellata  Rud.,  and 
consequently,  of  Taenia  percae  Miiller,  as  Kraemer,  Riggenbach,  Bene- 
dict, Railliet,  and  others  have  thought.  That  belief  was  founded  on  a 
mistaken  identification  by  Kraemer  (1892).  Since  Dujardin  (1845) 
there  have  been  but  few  records  of  examinations  of  Gasterosteus.  One 
of  these  records  is  by  Lonnberg,  another  by  Schneider.  They  examined 
Oasterosteus  pungitius,  while  the  type  host  of  T.  filicollis  is  Gasterosteus 


11]  PROTEOCEPHAUDAE  —  LA  RUE  11 

aculeaius  and  so  far  as  the  writer  can  discover  there  have  been  almost 
no  records  of  collection  of  parasites  from  that  host  since  Dujardin.  The 
diagnoses  of  Taenia  filicollis  much  resemble  those  of  Taenia  ambigua. 
Dujardin  however  evidently  considered  these  two  species  as  different 
for  he  records  both.  At  the  present  time  but  a  single  species  is  well 
known  from  Gasterosteous,  viz,  Proteocephalus  ambiguus  (Dujardin). 
yyThe  writer  considers  however  that  P.  ambiguus  is  identical  with  P. 
filicollis  (Rud.)  and  hence  the  genera  Proteocephalus  Weinland  and 
Ichthyotaenia  Lonnberg  being  based  on  the  same  species  are  synonyms 
and  the  earlier  name  should  be  retained. 

The  synonymy  of  the  name  Tetracotylus  Monticelli  (1891)  has  not 
yet  been  discussed.  This  name  was  based  on  the  description  of  Taenia 
coryphicephala  altho  Monticelli  failed  to  designate  it  as  the  type  species. 
Braun  (1894-1900)  stated  that  this  genus  was  based  on  T.  coryphiceph- 
ala and  this  action  was  considered  by  Hall  (1910)  to  be  tantamount 
to  the  designation  of  a  type.  Braun  regards  Tetracotylus  to  be  a  syno- 
nym of  Ichthyotaenia  which  makes  it  a  synonym  of  Proteocephalus  pro- 
vided that  the  type  of  Tetracotylus  belongs  in  the  same  genus  with 
Taenia  filicollis  (Rud.),  the  type  of  Proteocephalus.  That  Tetracotylus 
coryphicephala  does  not  belong  to  the  same  genus  with  Proteocephalus 
filicollis  has  been  shown  by  the  writer  in  the  descriptive  study  of  the 
former  species  (vide  infra).  Moreover  the  writer  has  shown  that  this 
form  does  not  belong  with  any  genus  of  cestodes  at  present  known.  It 
is  then  necessary  to  examine  into  the  status  of  the  name  Tetracotylus  as 
a  generic  name  for  Taenia  coryphicephala. 

The  availability  of  Tetracotylus  has  been  questioned  because  of  the 
name  Tetracotyle  Filippi  (1854),  These  two  names  are  not  spelled  alike 
and  are  therefore  not  homonyms,  Braun  (1894-1900)  suggests  that 
Tetracotylus  has  been  used  for  Tetracotyle.  If  such  improper  use  has 
been  made  of  the  former  name  prior  to  the  time  when  Monticelli  pro- 
posed it  then  Tetracotylus  has  been  rendered  unavailable.  The  writer 
has  not  been  able  to  find  evidence  of  such  improper  use,  yet  because  of 

^Braun's  statement  of  the  fact  and  his  objection  to  the  use  of  the  name 
^Xwthe  writer  has  proposed  the  name  Monticellia  in  honor  of  Professor  Mon- 

<  ticelli  who  has  done  so  much  for  our  knowledge  of  this  group,  as  a  name 
to  take  the  place  of  the  invalidated  Tetracotylus,  The  type  of  this 
genus  is  Monticellia  coryphicephala  (Monticelli), 

The  genus  is  to  be  defined  thus :  Monticellia  La  Rue :  Head 
small,  globose,  without  folds  or  lappets  of  tissue  encircling  suckers. 
Suckers  sessile  and  without  accessory  areola.  No  rostellum.  Testes, 
vitellaria  and  uterus  entirely  outside  of  the  inner  longitudinal  muscle- 
sheath.     Vitellaria  composed  of  scattered  follicles  which  form  broad 


12  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [12 

lateral  fields.  Testes  numerous,  forming  a  single  broad  dorsal  field  be- 
tween vitellaria.  Uterus  ventral,  with  many  lateral  pouches.  Genital 
pore  marginal,  irregularly  alternating.  Ovary  bilobed  and  situated 
partly  within  and  partly  outside  the  inner  muscle-sheath.  Sexual  organs 
in  general  as  in  Proteocephalidae.  In  Siluridae.  The  type  of  the  genus 
is  Monticellia  coryphicephala  {Tetracotylus  coryphicephala  Monticelli). 

The  position  of  this  genus  in  the  order  Tetraphyllidea  is  difficult  to 
determine.  It  apparently  does  not  belong  to  any  of  the  families  as  now 
defined  therefore  the  writer  suggests  for  it  the  family  name  Monticelli- 
dae  with  the  following  characters:  Head  small.  Suckers  sessile  and 
without  accessory  areola.  Internal  anatomy  as  in  the  type  genus  Monti- 
cellia. 

There  still  remains  the  question  as  to  whether  the  name  Proteo- 
cephalus  is  available.  That  name  has  been  objected  to  on  account  of  the 
name  Proteocephala  suggested  by  de  Blainville  (1828)  for  a  cestode 
family.  These  two  names  are  not  homonyms  and  it  is  agreed  that  the 
use  of  a  name  to  designate  a  family  does  not  invalidate  it  for  use  subse- 
quently as  a  generic  name.  The  writer  therefore  retains  the  name 
Proteocephalus  Weinland  as  the  generic  designation  for  the  genus  of 
which  Taenia  filicollis  Rud.  is  the  type. 

Having  determined  that  Monticellia  coryphicephala  (Monticelli) 
does  not  belong  in  the  family  Proteocephalidae  because  of  its  peculiar 
organization,  it  is  necessary  to  redefine  that  family  thus:  Family  Pro- 
teocephalidae: Heads  small.  Suckers  sessile  and  without  accessory 
areola.  Fifth  sucker  functional,  vestigial,  or  lacking.  No  rostellum. 
Genital  organs  in  general  as  in  other  Tetraphyllideans.  Genital  pores 
marginal,  irregularly  alternating.  Vitellaria  lateral,  follicular,  follicles 
closely  grouped  about  a  central  conducting  tubule.  Ovary  bilobed, 
posterior.  Oocapt,  ootype,  shell  gland,  uterine  passage  present.  Uterus 
with  lateral  outpocketings  and  one  or  more  preformed  ventral  uterine 
openings.  Vitellaria,  testes,  ovary  and  uterus  within  the  inner  longi- 
tudinal muscle-sheath. 

Habitat:    In  fresh-water  fish,  amphibia,  and  aquatic  reptiles. 

In  this  family  belong  the  following  genera  which  are  here  defined: 

(1)     Proteocephalus  Weinland  1858 

"With  characters  of  the  family.  Head  globose  or  conical,  flattened 
dorsoventrally.  No  rostellum.  No  spines  or  hooks.  No  fold  of  tissue 
encircling  base  of  head  or  enfolding  suckers.  Suckers  circular  or  oval. 
Fifth  sucker  functional  or  vestigial,  rarely  lacking.  Testes  in  a  broad 
field  between  vitellaria.  Parenchyma  with  close  meshes.  Musculature 
well  developed.  Eggs  with  three  membranes.  Habitat :  In  fresh-water 
Ml. 


13]  PROTEOCEPHAUDAE  —  LA  RUE  13 

Type  species:    Proteocephalus  filicolUs  (Rudolphi) 
Syn. :    Proteocephalus  amhiguus  (Dujardin) 

(2)  Choanoscolex  La  Rue  1911 

With  characters  of  family.  Head  conical  with  a  fold  of  tissue  at 
the  base  partially  covering  suckers.  No  rostellum,  no  hooks,  no  spines. 
Oenital  organs  as  in  Proteocephalus.    Habitat :  In  Siluridae. 

Type  species:     Choanoscolex  ahscisus  (Riggenbach). 

(3)  Corallohothrium  Fritsch  1886 

"With  characters  of  family.  Scolex  with  four  suckers  situated  on 
the  flat  anterior  face  of  the  head.  Many  irregular  folds  and  lappets  of 
tissue  about  margin  of  anterior  surface;  may  enclose  suckers  as  in  a 
corolla.  No  rostellum.  No  hooks  nor  spines.  Neck  broad,  short.  Habi- 
tat :    In  Siluridae. 

Type  species:    Corallohothrium  solidum  Fritsch. 

(4)  Crepidohothrium  Monticelli  1899 

With  characters  of  family.  Head  large,  swollen,  pjrramidal,  tetrag- 
onal, unarmed.  Suckers  large,  inversely  cordate,  posterior  margin  in- 
terrupted and  re-entrant  into  sucker  cavity.  Fifth  sucker  vestigial. 
Oenital  apertures  marginal,  irregularly  alternating.  Vaginal  opening 
dorsal  to  cirrus  pouch.  Vagina  anterior  or  posterior  to  cirrus-pouch. 
Testes  in  two  lateral  fields  anterior  to  ovary.  Habitat:  In  Boidae, 
South  America. 

Type  species:     Crepidohothrium  gerrardii  (Baird) 

(5)  Acanthotaenia  von  Linstow   1903 

With  characters  of  family.  Scolex  rather  small.  With  four  rounded 
suckers.  Apex  of  head  conical  in  which  may  be  situated  a  vestigial  fifth 
sucker.  No  rostellum.  Cuticula  of  head  and  anterior  part  of  body  cov- 
ered with  minute  spines  or  bristles.  Segmentation  indistinct.  No  over- 
lapping of  segments.  Testes  in  two  lateral  fields  anterior  to  ovary. 
Ovarian  lobes  may  be  branched.  Vagina  anterior  or  posterior  to  cirrus. 
Habitat:    In  Varanidae  and  Hylidae. 

Type  species :    Acanthotaenia  shipleyi  von  Linstow 

(6)  Ophiotaenia  La  Rue  1911 

With  characters  of  family.  Head  globose  or  somewhat  tetragonal. 
No  rostellum.  No  hooks  or  spines.  Suckers  circular  or  oval,  with  mar- 
gins entire.  Fifth  sucker  vestigial.  Neck  usually  long.  Testes  in  two 
long  lateral  fields  anterior  to  ovary.  Vagina  anterior  or  posterior  to 
cirrus-pouch.     Ovary  bilobed,  flattened,  sometimes  alate.     Parenchyma 


V 


14  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [14- 

with  fine  meshes.  Musculature  weak.    Habitat:  In  aquatic  snakes,  Cro- 
talinae,  Colubridae,  Elapinae  and  Amphibia. 

Tyj>e  species :    Ophiotaenia  perspicua  La  Rue  1911. 

In  the  descriptive  section  the  greater  number  of  species  of  the 
above  named  genera  have  been  described,  with  exception  of  species  of 
Acanthotaenia  and  Corallobothrium.  It  has  been  impossible  to  write  up 
the  descriptions  of  certain  species  inquirendae  of  which  no  material 
could  be  obtained,  and  for  which  only  very  meager  descriptions  have 
ever  been  written.  It  is  hoped  that  these  may  be  properly  considered 
at  a  later  time. 


METHODS  OF  TECHNIQUE 

The  following  methods  have  been  used  by  the  writer  in  the  work 
on  this  group  of  cestodes.  To  a  large  extent  they  may  be  used  with 
success  on  all  groups  of  cestodes  although  it  should  be  understood  that 
certain  methods  which  give  admirable  results  with  the  relatively  small 
and  thin  cestodes  here  dealt  with  will  not  give  equally  good  results  if 
used  on  the  large  forms  such  as  Taenia. 

The  larger  forms  were  picked  out  of  the  intestinal  contents,  care 
being  taken  to  free  the  head  if  the  worm  was  attached  to  the  mucosa. 
These  were  then  repeatedly  dipped  in  the  killing  solution  until  the  worm 
ceased  to  contract.  The  worm  was  allowed  to  lie  for  15  minutes  to  2 
hours  in  the  same  fluid.  Metallic  instruments  are  to  be  avoided  if  cor- 
rosive sublimate  solutions  are  used  for  fixation.  When  the  smaller 
worms  were  encountered  the  whole  intestine  slit  open  was  placed  in  a 
small  quantity  of  physiological  saline  solution  in  a  bottle  which  was  then 
shaken  vigorously  for  about  3  minutes,  the  killing  fluid  was  added  and 
the  whole  then  shaken  for  one  half  minute.  This  is  according  to  the 
method  of  Looss.    The  fixative  was  permitted  to  act  3  to  10  hours. 

The  killing  fluids  used  were  hot  5%  solution  of  formaldehyde,  and 
hot  or  cold  saturated  aqueous  solution  of  corrosive  sublimate  to  which 
was  added  glacial  acetic  acid  to  make  1  to  2%.  Some  other  fluids  were 
tried  but  nothing  gave  better  results  for  the  purposes  of  this  study  than 
the  corrosive  acetic  mixture  used  hot  or  cold.  For  most  of  the  worms 
the  cold  solution  was  preferable  to  the  hot  which  sometimes  gave  rise  to 
artifacts  if  used  at  too  high  a  temperature.  In  no  cases  were  the  worms 
stupefied  before  killing. 

The  usual  methods  were  used  for  hardening  and  dehydrating.  Speci- 
mens were  usually  preserved  in  85%  alcohol  after  running  up  through. 


15]  PR0TE0CEPHAL1DAE~LA  RUE  15 

the  grades.  Sometimes  after  the  corrosive  acetic  fixation  5%  formalin 
was  used  for  a  preservative  with  uniformly  excellent  results. 

Sections  were  cut  5  to  10  micra  thick  for  the  study  of  histological 
detail  and  20  micra  when  grosser  morphological  details  were  sought. 
The  sections  were  stained  with  haematoxylin  mixtures,  either  Delafield's 
haematoxylin  or  Mayer's  haemalum,  and  decolorized  in  the  manner 
approved  for  these  stains.  Methods  of  staining  in  toto  followed  by  sec- 
tioning were  used  with  great  success  at  times.  For  this  purpose  Ehr- 
lich's  acid  haematoxylin  much  diluted  with  50%  alcohol  gave  the  best 
results.  For  a  contrast  stain  eosin  in  95%  alcohol  was  used  on  the  sec- 
tions.   Acid  fuchsin  also  in  95%  alcohol  was  sometimes  used  effectively. 

Preparations  in  toto  were  much  used  and  were  found  to  be  of  great 
value  in  mapping  out  the  relationships  of  the  organs  of  the  proglottids. 
Frequently  these  methods  showed  everything  to  be  desired  except  the 
histology  of  the  organs.  In  some  cases  even  histological  details  were 
well  revealed  by  these  methods.  The  stains  which  were  tried  for  staining 
in  toto  were  Mayer's  paracarmine,  Grenacher's  borax  carmine,  some 
alcoholic  cochineal  mixtures,  Mayer's  haemalum,  Delafield's  haematoxy- 
lin, and  Ehrlich's  acid  haematoxylin.  None  of  the  carmine  or  cochineal 
stains  were  very  successful  for  none  of  them  show  the  boundaries  of 
cestode  structures  sharply.  The  parenchyma  in  which  the  genital  organs 
lie  always  retained  too  great  an  amount  of  these  stains  to  permit  a  clear 
view  of  the  genital  organs  themselves.  The  haematoxylins,  however, 
usually  gave  wonderfully  clear,  sharp  pictures  of  the  genital  organs. 
It  was  at  times  possible  to  work  out  such  minute  structures  as  vasa 
efferentia  almost  in  their  entirety  from  such  preparations  in  toto.  The 
three  haematoxylin  stains  were  found  to  be  about  equally  good. 

In  using  these  stains  it  was  the  practice  to  dilute  the  stain  with  the 
proper  diluent.  Relatively  large  quantities  of  the  diluted  stain  were 
used  for  each  lot  of  material.  The  stain  was  permitted  to  act  over  night 
(10  to  15  hours)  at  room  temperature.  The  excess  of  the  stain  was  then 
removed  by  washing  in  distilled  water  and  the  tissue  passed  through  the 
grades  of  alcohol  to  70%  where  it  was  decolorized  rapidly  by  adding 
hydrochloric  acid  to  make  a  0.5  to  1.0%  solution.  The  object  was  to 
remove  the  stain  from  the  peripheral  tissues  at  a  rapid  rate  and  mean- 
while leave  the  stain  in  the  deeper  lying  tissues.  In  this  method  the 
duration  of  the  acid  bath  is  usually  short  depending  upon  the  size  of  the 
piece  and  the  character  of  the  stain  taken  by  the  tissue,  and  upon  the 
character  of  the  tissue  itself.  In  general  it  is  desirable  to  decolorize 
until  a  light  reddish  blue  stain  remains  and  until  many  of  the  internal 
structures  can  be  distinguished  while  the  tissue  is  still  in  the  alcoholic 
medium.     When  in  the  judgment  of  the  operator  the  proper  stain  is 


16  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [16 

attained  the  tissues  are  placed  in  neutral  alcohol  and  then  into  70% 
alcohol  rendered  slightly  alkaline  by  the  addition  of  a  few  drops  of  an 
aqueous  solution  of  sodium  carbonate. 

Preparations  were  not  flattened  but  were  straightened  out  on  a  slide 
and  over  this  was  placed  another  slide  which  was  supported  by  strips 
of  paper  of  such  a  thickness  that  little  or  no  pressure  was  exerted  on 
the  specimen  by  the  slides.  Dehydration  and  clearing  were  accomplished 
while  the  preparation  was  thus  kept  straight.  Xylol  and  cedarwood  oil 
were  used  as  clearing  agents.    Preparations  were  mounted  in  balsam. 

The  methods  outlined  above  yielded  very  satisfactory  preparations 
for  the  study  of  these  cestodes  and  they  have  also  been  used  by  the 
writer  on  other  cestodes  and  on  trematodes  with  great  success.  It  is 
noteworthy  that  the  carmine  stains  give  beautiful  preparations  of  trem- 
atodes in  toto  but  fail  almost  entirely  for  cestodes.  For  the  cestodes 
these  stains  fail  because  they  do  not  sharply  and  clearly  outline  the 
sexual  organs  as  they  do  in  trematodes  though  not  better  than  do  the 
haematoxylins.  In  the  judgment  of  the  writer  the  use  of  the  carmine 
stains  on  cestode  material  has  been  responsible  for  many  errors  in  the 
interpretation  of  cestode  structures. 


ANATOMY  AND  HISTOLOGY  OF  PROTEOCEPHALIDS 

This  section  deals  in  a  very  general  way  with  the  anatomy  and 
histology  of  the  Proteoeephalidae,  placing  emphasis  on  the  usual  char- 
acter of  the  structures  encountered  and  at  times  calling  attention  to 
variations  in  this  general  plan.  In  the  descriptive  part  of  the  work  the 
ordinary  histological  details  and  many  of  the  anatomical  details  of  lesser 
importance  have  been  omitted  or  mentioned  in  only  a  casual  way. 
"Whenever  the  character  of  a  structure  has  departed  from  the  usual  the 
fact  has  been  noted  in  more  or  less  detail. 

The  cestodes  of  the  family  are  constructed  on  a  uniform  plan  which 
has  been  thoroly  discussed  by  Monticelli  (1891),  Kraemer  (1892),  Rig- 
genbach  (1896),  Benedict  (1900),  Schneider  (1905),  Schwarz  (1908), 
and  La  Rue  (1909).  Some  new  points  are  brought  out  in  the  present 
paper.  The  histological  structures  of  these  cestodes  have  been  pretty 
well  worked  out  prior  to  the  present  time.  Von  Linstow  ( 1891 ) ,  Monti- 
celli (1891),  Kraemer  (1892),  and  Riggenbach  (1896)  did  pioneer  work 
on  the  histology  of  the  group.  Riggenbach  (1896)  and  Benedict  (1900) 
made  very  careful  and  accurate  studies  of  the  histology  of  certain  spe- 
cies of  fish  Proteocephalids  and  were  able  to  clear  up  some  of  the  earlier 
misconceptions.     No  careful  work  on  the  histology  of  the  amphibian 


17]  PROTEOCEPHALIDAE  —  LA  RUE  17 

Proteocephalids  was  made  until  the  work  of  the  writer  (La  Rue,  1909) 
on  Ophiotaenia  (Proteocephalus)  filaroides.  Subsequent  work  on  the 
histology  of  the  eestodes  of  snakes  has  shown  but  little  new,  altho  it 
must  be  admitted  that  only  little  has  been  done  on  the  group. 

The  scolex  is  usually  of  small  size  and  rather  inconspicuous.  Among 
the  fish  Proteocephalids  the  heads  range  in  size  from  a  little  more  than 
0.1  mm.  in  breadth  to  about  1.  mm.,  tho  in  at  least  one  species,  P.  sulca- 
ius,  the  breadth  of  this  region  attains  as  much  as  1.75  mm.  Among  the 
amphibian  and  reptilian  Proteocephalids  the  heads  range  from  about 
0.2  mm.  to  1,75  mm.  in  breadth.  In  general  the  head  is  not  sharply 
delimited  from  the  neck,  which  may  be  of  nearly  equal  width  or  in 
exceptional  cases  even  wider  for  a  short  distance  than  the  head.  In 
form  the  heads  show  considerable  variation.  They  are  usually  more  or 
less  globose  or  conical,  and  somewhat  flattened  dorsoventrally.  In  a 
few  cases  the  head  inay  present  a  more  or  less  flattened  tetragonal 
anterior  face.  The  surface  of  the  heads  may  be  smooth  or  marked  by 
shallow  or  deep  grooves  between  the  suckers.  As  a  rule  the  heads  have 
no  folds  or  lappets  of  tissue  about  the  suckers  or  on  the  anterior  face. 
Such  structures  are  found,  however,  in  Corallobothrium  and  to  a  more 
limited  extent  in  Choanoscolex.  The  special  characters  of  the  heads  of 
each  species  are  taken  up  in  the  descriptive  part  of  the  work.  The 
heads  of  Proteocephalidae  and  Monticellidae  differ  from  those  of  other 
families  of  Tetraphyllidea  in  that  the  suckers  are  sessile,  and  have  no 
accessory  areola.  Even  when  the  head  is  deeply  furrowed  the  sucker 
never  has  the  appearance  of  having  a  stalk.  Suckers  may  be  prominent 
or  inconspicuous  and  so  may  influence  the  general  appearance  of  the 
head. 

A  rostellum  is  unknown  among  the  species  of  this  family,  altho 
certain  species  have  been  erroneously  reported  to  have  such  an  organ. 
It  is  possible,  tho  not  probable,  that  some  species  of  Acanthotaenia  have 
a  rostellum.  Some  of  these  species  have  long  apical  prominences  on  the 
head,  but  except  for  the  elongation  of  this  apex  and  the  presence  of 
cuticular  spines  there  is  no  evidence  for  considering  this  structure  to  be 
a  rostellum.  The  spines  are  not  hooks  but  are  cuticular  structures  which 
may  extend  over  the  anterior  part  of  the  body.  Neither  elongated  tip 
nor  cuticular  spines  are  good  criteria  of  the  presence  of  a  rostellum. 
To  determine  the  presence  of  a  rostellum  one  should  look  to  the  inner 
structure  of  the  apex,  its  musculature  and  protrusibility,  and  to  the 
origin  of  this  rostellum-like  tip.  In  the  tip  of  the  heads  of  many  species 
which  do  not  possess  a  functional  fifth  sucker  there  has  been  found  a 
peculiar  structure  consisting  of  massed  cells  and  sometimes  a  few  mus- 
cular elements.    This  structure  has  been  considered  to  be  a  rudimentary 


18  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [18 

rostellum.  Such  a  supposition,  however,  is  incorrect,  for  in  its  devel- 
opment it  is  a  fifth  sucker.  A  more  complete  discussion  of  this  rudi- 
mentary or  vestigial  fifth  sucker  is  given  under  the  proper  caption 
(vide  infra).  ' 

All  species  are  provided  with  four  suckers  and  some  with  an  apical 
fifth  sucker  in  addition.  The  latter  may  be  either  functional  or  vesti- 
gial. The  four  suckers  are  sessile,  prominent,  or  inconspicuous  cup- 
shaped  organs  which  are  usually  situated  on  the  broadest  part  of  the 
head  or  just  anterior  to  the  broadest  zone.  In  Corallobothrium  the 
suckers  are  situated  on  the  flattened  anterior  face  of  the  head.  The 
suckers  do  not  possess  accessory  areola  nor  are  they  provided  with  hooks 
in  their  cavities  or  on  their  margins.  Acanthotaenia  may  have  cuticular 
spines  within  the  sucker  cavity.  Suckers  are  usually  rounded  or  oval 
in  outline  and  may  have  deep  cavities.  The  margin  is  usually  entire 
but  is  at  times  interrupted.  Some  peculiar  cases  occur :  Proteocephalus 
Cyclops  has  suckers  pointed  at  the  posterior  end.  The  suckers  of 
P.  singularis  (Fig.  25)  are  flat,  thin  and  weakly  muscled  and  have  a 
peculiar  upturned  point  at  the  anterior  margin.  The  deep  grooves  be- 
tween the  suckers  of  this  species  cause  these  organs  to  stand  out  promi- 
nently. Crepidobothrium  gerrardii  (Figs.  12,  13,  33,  34,  123,  and  124) 
likewise  has  peculiar  suckers  in  that  each  has  an  interrupted  lower 
margin  which  forms  a  point  re-entrant  into  the  sucker  cavity.  Deep 
grooves  between  the  suckers  give  the  head  a  lobed  appearance  and  cause 
the  suckers  to  be  prominent.  Proteocephalus  osculatus  (Fig.  162)  has 
a  fifth  sucker  which  is  said  to  be  covered  with  minute  spinelets.  The 
suckers  of  some  of  the  Acanthotaenia  are  covered  with  minute  spinelets. 
It  is  deemed  unnecessary  to  enter  into  a  discussion  of  the  histologj"  of  the 
suckers  of  this  group.  Benedict  (1900),  Kraemer  (1892),  and  La  Rue 
(1909)  have  discussed  this  subject  in  detail. 

A  functional  fifth  sucker  situated  at  the  apex  of  the  head  is  present 
in  many  species  of  Proteocephalus  but  is  not  known  among  the  other 
genera  of  the  family.  Other  species  of  Proteocephalus,  Ophiotaenia, 
Crepidobothrium,  and  Acanthotaenia  are  known  to  possess  a  structure 
which  the  writer  (1909)  called  an  end-organ  and  which  Johnston  (1909 
et  seq.)  has  called  an  apical  muscle-plugy^ther  species  of  Proteocepha- 
lus and  all  known  species  of  Corallobothrium  and  Choanoscolex  do  not 
possess  this  structure  or  the  functional  fifth  sucker  while  some  species 
of  Proteocephalus  and  Ophiotaenia  have  not  been  investigated  for  it. 
The  writer  has  now  determined  that  this  organ  is  a  vestigial  fifth  sucker. 
The  fifth  sucker  when  functional  is  usually  smaller  than  the  others  and 
it  possesses  all  the  histological  structures  of  other  Proteocephalid  suckers. 
The  basement  membrane  and  the  muscles  have  the  same  relations  as  in 
other  suckers. 


19]  PROTEOCEPHALIDAE  —  LA  RUE  19 

The  vestigial  fifth  sucker  in  the  adult  head  is  represented  by  a  mass 
of  cells  or  of  nuclei  and  at  times  a  few  fibers  surrounded  by  a  basement 
membrane.  It  is  entirely  sunken  into  the  tissues  of  the  head  and  has 
lost  all  connection  with  the  exterior.  An  examination  of  this  structure 
in  an  adult  head  yields  no  clue  to  its  probable  origin,  but  if  heads  of 
plerocercoids  of  a  species  which  has  this  structure  in  the  adult  be  exam- 
ined it  is  seen  to  be  a  degenerate  fifth  sucker.  In  the  plerocercoid  this 
degenerating  fifth  sucker  has  a  sucker  cavity  communicating  with  the 
exterior.  It  has  basement  membranes  and  a  full  complement  of  muscles 
altho  some  of  the  muscles  may  be  somewhat  displaced  and  undergoing 
atrophy.  The  origin  of  this  structure  has  been  worked  out  in  some 
detail  for  Ophiotaenia  filaroides  and  the  results  are  given  in  the  descrip- 
tion of  that  species.  Its  structure  is  illustrated  by  drawings  which  are 
reproduced  here  (Figs.  27,  28).  The  structure  of  the  adult  organ  was 
given  in  the  writer's  paper  on  this  cestode  (La  Rue  1909). 

A  vestigial  fifth  sucker  which  has  not  reached  such  a  state  of 
degeneracy  as  is  found  in  many  of  the  species  occurs  in  the  head  of 
Crepidohothrium  gerrardii.  In  this  species  the  sucker  tho  so  greatly 
reduced  in  size  as  to  be  overlooked  in  an  examination  of  toto  prepara- 
tions still  retains  its  cavity  which  is  yet  in  connection  with  the  exterior 
through  a  small  pore.  The  sucker  is  surrounded  by  a  basement  membrane 
and  it  shows  some  traces  of  muscles  altho  the  material  studied  was  too 
poor  to  make  a  good  histological  study.  A  more  complete  description 
of  this  vestigial  sucker  is  to  be  found  in  the  description  of  Crepidohoth- 
rium gerrardii.  Its  structure  is  delineated  in  figure  42.  In  his  descrip- 
tion of  these  vestigial  structures  found  in  O.  filaroides  and  in  C.  gerrar- 
dii the  writer  has  conclusively  shown  that  the  organ  under  discussion  is 
a  vestigial  fifth  sucker  and  not  a  vestigial  or  rudimentary  rostellum.  It 
seems  highly  probable  that  similar  structures  found  in  other  species  of 
the  group  will  prove  to  be  vestigial  fifth  suckers. 

The  writer  can  see  no  reason  for  assuming  that  this  structure  is  a 
rudimentary  rostellum.  So  far  as  known  a  well  developed  rostellum 
does  not  occur  among  the  Tetraphyllidea  but  is  characteristic  of  many 
species  of  Cyclophyllidea.  Unless  an  apical  sucker  is  a  transitional  stage 
in  the  development  (phylogenetie)  of  a  rostellum  the  writer  sees  no 
reason  for  regarding  this  vestigial  fifth  sucker  as  a  rudimentary  rostel- 
lum. The  knowledge  concerning  this  structure  is  incomplete  and  studies 
of  developmental  stages  of  Proteocephalid  species  possessing  vestigial  or 
functional  fifth  suckers  should  be  made  in  comparison  with  similar  de- 
velopmental stages  of  species  of  Calliobothriura  and  Anthocephalus 
which  according  to  Lang  (1881)  and  Monticelli  (1888)  have  certain 
rudimentary  structures  that  seem  to  be  like  the  oral  suckers  of  trema- 


20  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [20 

todes.  Likewise  a  study  should  be  made  of  certain  species  of  Cyclophyl- 
lidea  which  are  known  to  possess  a  rudimentary  or  a  vestigial  rostellum. 
No  definite  solution  of  this  problem  can  be  reached  without  such  a  com- 
parative study.  Data  as  to  the  presence  or  absence  of  the  fifth  sucker 
in  species  of  the  group,  whether  functional  or  vestigial,  is  to  be  found 
in  the  tables  of  the  principal  characters  of  the  cestodes  of  this  family. 

The  nervous  system  is  made  up  of  a  nerve  ring  which  occurs  in  the 
neighborhood  of  the  suckers  and  one  pair  of  main  lateral  nerve  trunks 
in  the  strobila.  As  described  by  the  writer  (1909)  for  Ophiotaenia  fila- 
roides  the  nerve  ring  is  somewhat  octagonal  in  shape  and  at  its  corners 
nerve  processes  extend  out  to  the  suckers.  From  the  ring  main  trunks 
extend  back  into  the  strobila.  In  the  strobila  a  single  pair  of  main 
lateral  trunks  may  be  seen  just  within  the  lateral  fields  of  the  muscle 
sheath.  No  accessory  nerve  trunks  have  been  made  out  by  the  writer. 
In  Ophiotaenia  filaroides  the  nerve  trunk  passes  dorsal  to  the  cirrus  and 
the  vagina. 

In  all  species  the  neck  is  a  more  or  less  poorly  defined  unsegmented 
region  between  the  head  and  the  segmented  part  of  the  strobila.  This 
region  may  be  several  millimeters  long  in  certain  species  while  in  others 
it  is  said  to  be  entirely  lacking  or  may  be  no  more  than  0.3  to  0.5  mm. 
long.  It  is  usually  narrower  than  the  scolex  tho  in  exceptional  cases  in 
which  the  neck  muscles  may  be  unduly  contracted  the  neck  is  broader 
than  the  scolex.  Much  confusion  has  arisen  over  the  statements  of  the 
older  investigators  who  described  the  neck  of  certain  species  as  being  long 
or  very  long  without,  however,  giving  measurements.  These  adjectives 
are  relative  terms.  If  the  cestode  is  small,  0.5  to  1  mm.  broad  and  20  to 
100  mm.  in  length,  a  neck  which  is  1  to  5  mm.  long  seems  long  or  very 
long.  A  neck  of  that  length  on  a  cestode  as  large  as  Crepidohothrium 
gerrardii  which  is  several  millimeters  broad  and  many  centimeters  long 
would  be  called  short.  In  this  species  an  unsegmented  region  0.5-1  mm. 
long  is  ignored  by  some  authors  and  the  specimen  is  reported  as  having 
no  neck.  It  is  essential  that  actual  measurements  of  the  unsegmented 
region  be  made.  For  this  purpose  stained  specimens  are  necessary  be- 
cause unstained  specimens  will  not  show  segmentation  plainly  and  also 
because  external  wrinkles  or  folds  may  simulate  segmentation.  In  re- 
cording measurements  of  the  neck  the  writer  takes  the  length  from  the 
point  of  narrowing  behind  the  suckers  to  the  first  evident  traces  of  in- 
ternal segmentation.  In  a  few  species  another  system  has  been  found 
necessary  but  in  these  cases  the  method  of  measuring  has  been  stated. 
The  breadth  given  is  the  narrowest  place  in  the  unsegmented  region. 

The  segmented  part  of  Proteocephalids  varies  greatly  in  length, 
breadth,  and  thickness.    In  some  species  the  strobila  is  small,  being  not 


21]  PROTEOCEPHALIDAE  —  LA  RUE  21 

more  than  a  few  millimeters  long  and  less  than  1  mm.  broad,  while  in 
others  it  may  attain  a  length  of  60  or  more  centimeters  and  an  extreme 
breadth  of  2  to  4  mm.  Such  extreme  size  is  infrequent  even  for  the  large 
species.  Many  of  the  species  of  Proteocephalus  are  thick  and  fleshy 
while  a  few  of  them  are  thin  and  flat.  So  also  are  the  known  species  of 
the  family  Monticellidae.  Segmentation  may  be  evident  or  obscure. 
Perhaps  in  no  species  is  it  as  evident  as  it  is  in  some  species  of  Taenia. 
The  proglottids  are  usually  without  sharp  posterior  angles  and  they  are 
attached  along  their  entire  width.  Transverse  intersegmental  furrows 
if  present  are  shallow. 

The  youngest  proglottids  are  very  indistinct  and  are  distinguished 
in  stained  preparations  as  faint  cross  bands  of  alternate  dark  and  light. 
These  are  darkly  stained  areas  of  rapidly  differentiating  parenchyma 
alternating  with  the  unstained  future  septa.  Almost  without  exception 
the  young  proglottids  are  much  broader  than  long.  The  reverse  of  this 
is  true  only  when  the  anterior  part  of  the  body  is  in  an  extremely  attenu- 
ated condition.  As  the  young  proglottids  develop  they  increase  in 
length  and  breadth,  tho  the  length  usually  increases  more  rapidly  than 
the  breadth.  The  transition  from  young  to  mature  proglottids  is  grad- 
ual. Mature  proglottids  are  those  in  which  the  sexual  organs  have  at- 
tained maturity  and  egg  production  is  about  to  begin  or  may  just  have 
begun.  Such  proglottids  are  usually  broader  than  long  or  quadrate. 
The  length  is  much  greater  than  in  the  young  proglottids.  Mature  prog- 
lottids are  the  best  ones  in  which  to  study  the  greater  part  of  the  genital 
system  tho  of  course  a  study  of  the  uterus  must  be  made  from  the  ripe 
proglottids. 

Those  proglottids  in  which  egg  production  is  well  along  or  complete 
are  spoken  of  as  ripe  proglottids.  The  uterine  pouches  are  full  of  eggs 
and  the  ventral  openings  are  preparing  for  their  discharge.  Ripe 
proglottids  are  frequently  longer  than  broad  tho  the  reverse  is  not  un- 
common. They  are  normally  considerably  larger  than  mature  proglot- 
tids. In  them  the  vitellaria  and  testes  may  be  greatly  reduced  in  size  or 
almost  obliterated  by  the  great  mass  of  eggs  in  the  uterine  pouches.  An 
end-proglottid  is  frequently  present.  This  should  be  defined  as  the  first 
proglottid  to  be  differentiated  from  the  growing  zone  of  the  neck.  It 
still  retains  certain  of  the  characters  of  the  posterior  end  of  the  plero- 
cercus  which  is  not  lost  in  the  passage  of  the  young  tapeworm  through  the 
stomach  of  the  host.  It  is  pointed  posteriorly  and  has  at  the  tip  a 
median  excretory  pore  through  which  the  excretory  products  are  dis- 
charged. Just  anterior  to  this  pore,  in  some  species,  is  a  contractile 
bladder.  In  the  species  examined  by  the  writer  a  number  of  end-proglot- 
tids  have  been  found;  some  of  these  contained  eggs  and  so  are  to  be 


22  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [22 

considered  as  functional.  Since  ripe  proglottids  are  shed  after  or  about 
the  time  that  the  eggs  are  ripe  one  frequently  encounters  strobilas  which 
lack  the  end  proglottid.  The  loss  of  this  proglottid  is  not  to  be  consid- 
ered as  a  special  character,  for  it  is  common  to  many  eestodes  outside 
the  group. 

Two  pairs,  dorsal  and  ventral,  of  main  lateral  excretory  trunks  ex- 
tend through  the  strobila.  Of  these  the  ventral  vessels  are  the  larger. 
Some  additional  longitudinal  vessels  have  been  described  by  von  Linstow 
(1891)  in  Proteocephalus  longicollis.  It  seems  probable  that  he  saw  the 
cut  ends  of  branches  and  anastomoses.  Some  species  have  a  commissural 
vessel  connecting  the  ventral  excretory  vessels  at  the  posterior  end  of  the 
proglottid.  It  has  not  been  noted  in  other  species.  In  the  neck  region 
and  throughout  the  strobila  many  small  branches  arise  from  the  ventral 
vessels  and  some  also  from  the  dorsal  vessels  which  pass  to  the  surface 
of  the  worm  where  they  discharge  to  the  exterior  through  a  small  pore. 
These  openings  occur  more  fiV^uently  on  the  ventral  surface  but  may 
be  found  on  the  lateral  edga^  In  most  species  they  are  irregularly  dis- 
tributed but  in  Corallohothnum  lohosum  Riggenbach  (1896)  found  that 
they  regularly  came  to  the  exterior  at  the  posterior  comer  of  the  pro- 
glottid. This  finding  has  not  been  supported  by  subsequent  work  on 
other  species.  In  the  head  and  the  anterior  neck  region  there  are 
numerous  coils  of  vessels  and  anastomosing  branches  of  the  same  which 
extend  well  into  the  tip  of  the  head.  These  coils  lead  back  and  are  con- 
nected with  the  main  lateral  excretory  trunks.  At  the  posterior  end  of 
the  worm  the  excretory  vessels  discharge,  if  the  end-proglottid  is  pres- 
ent, through  a  common  excretory  pore  (Fig.  51).  Just  anterior  to  this 
pore  and  discharging  thi;ough  it  is  sometimes  a  pulsatile  bladder  which 
is  most  readily  seen  in  the  plerocercus.  The  flame  cells  at  the  ends  of 
minute  vessels  which  discharge  into  the  main  vessels  or  into  the  larger 
branches  are  situated  in  the  medullary  parenchyma  not  far  from  the 
main  excretory  vessels.  They  have  not  been  found  in  the  cortical  paren- 
chyma nor  in  the  mid-field  of  the  proglottid.  Their  distribution  in 
Ophiotaenia  filaroides  and  Proteocephalus  amhloplitis  has  been  thoroly 
worked  out  (La  Rue  1909). 

The  cuticula  and  the  subcuticular  structures  are  scarcely  worthy  of 
discussion  here  for  in  details  of  structure  they  do  not  differ  from  other 
eestodes.  In  the  Acanthotaenia  alone  are  there  special  features  of  the 
cuticula  to  which  attention  should  be  called.  Here  the  cuticula  is  thrown 
up  into  minute  cuticular  spines  which  are  too  minute  to  be  called  hooks 
or  booklets.  This  condition  is  most  prominent  on  the  head  and  neck 
but  may  also  occur  to  a  smaller  extent  over  the  cuticula  of  the  entire  body. 
These  features  have  been  emphasized  by  von  Linstow  (1903)  and  John- 


23]  PROTEOCEPHALIDAE  —  LA  RUE  23 

stou  (1909,  1911,  1912a).  Outside  of  this  genus  the  only  other  species 
to  have  cuticular  hooklets  or  spines  is  Proteocephalus  osculatus,  in  which 
they  are  reported  to  have  been  found  on  the  apical  fifth  sucker. 

The  parenchyma  resembles  that  of  other  cestodes  but  is  more  loose 
than  in  the  Cyclophyllidean  cestodes  which  the  author  has  examined. 
In  the  parenchyma  the  cell  outlines  are  indistinguishable  but  there  are 
numerous  structures  present  which  look  somewhat  like  minute  fibrillae. 
Large  fat  spaces  are  usually  visible  in  the  prepared  sections  and  some- 
times in  toto  preparations.  In  0.  filar oides  studied  by  the  author  (1909) 
the  fat  spaces  are  spheroidal  or  ovoidal  in  form  and  have  a  dimension  of 
0.03  to  0.045  mm.  Similar  spaces  have  been  observed  in  other  species  of 
the  family.  They  are  smaller  in  the  fish  cestodes  studied  and  larger  in 
the  snake  and  amphibian  cestodes.  The  fat  of  fresh  cestodes  or  of  those 
recently  killed  in  formol  is  readily  stained  by  osmic  acid  solutions  or  by 
an  alcoholic  solution  of  Sudan  III.  Other  specific  fat  stains  have  not 
been  tried.  The  character  of  the  parenchyma  affects  the  staining  prop- 
erties of  the  cestodes.  Those  cestodes  having  a  loose  parenchyma  make 
much  better  toto  preparations  than  do  those  which  have  the  dense 
parenchyma. 

In  general  the  parenchyma  of  the  group  of  snake  Proteocephalids 
is  a  more  open,  looser  network  than  that  of  the  fish  Proteocephalids. 
There  is  more  of  the  fibrillar  structure  present  in  the  parenchyma  of  the 
latter  than  of  the  former.  Whether  this  character  is  correlated  with 
differences  in  environmental  stimuli  is  not  known.  Perhaps  the  appear- 
ances of  the  parenchyma  of  cestodes  can  be  explained  as  are  the  appear- 
ances of  the  connective  tissues  of  higher  animals.  If  this  be  true  then 
it  is  evident  that  the  closer  woven  texture  of  the  parenchyma  in  fish 
cestodes  is  indicative  of  more  frequent  stresses  and  strains  placed  upon 
the  cestode  by  the  activity  of  the  host  or  by  the  movement  of  ingested 
materials  through  the  alimentary  tract. 

The  sub-cuticular  muscles  resemble  those  of  other  cestodes.  The 
circular  and  longitudinal  fibers  are  not  strong.  Within  the  layer  of  sub- 
cuticular cells  of  the  strobila  there  is  a  layer  of  longitudinal  muscle 
fibers  which  separates  the  cortical  from  the  medullary  parenchyma  in 
which  lie  almost  the  entire  generative  organs.  This  layer  of  fibers  forms 
what  is  known  as  the  inner  muscle  sheath.  The  sheath  is  well  developed 
in  some  species,  particuarly  in  the  larger  species  of  fish  Proteocephalids, 
while  in  the  weaker  species  and  in  Ophiotaenia  especially  it  is  poorly 
developed.  In  these  cases  the  muscles  are  few  in  number  and  separated 
from  each  other  by  intervals.  The  muscles  of  the  inner  sheath  pass 
through  the  intersegmental  septa.  Other  muscle  fibers  which  pass 
through  the  tissues  of  the  proglottid  from  dorsal  to  ventral  and  from 


24  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [24 

the  lateral  edges  are  frequently  encountered  but  are  not  as  numerous  as 
are  the  fibers  of  the  muscle  sheath;  they  do  not  occur  in  groups  but 
usually  singly.  In  the  region  of  the  septa  the  transverse  and  dorso- 
ventral  fibers  are  more  numerous  and  form  a  sort  of  muscular  plate. 

The  muscles  of  the  head  are  not  arranged  in  the  same  order  as  in 
the  Cyclophyllidea,  as  La  Rue  (1909)  pointed  out.  In  transverse  sections 
of  the  head  the  muscles  are  usually  encountered  as  follows:  Near  the 
tip  is  a  rhomboid  of  muscles,  deeper  down  a  transverse  muscle  cross  and 
then  a  diagonal  muscle  cross,  the  last  two  forming  a  muscle  star.  The 
muscle  star  occurs  in  the  region  of  the  suckers  but  in  certain  cases 
(Riggenbach  1896)  it  may  extend  below  the  level  of  the  suckers.  Usually 
as  soon  as  the  suckers  have  been  passed  the  muscles  take  on  an  arrange- 
ment typical  of  the  muscle  arrangement  in  the  neck  and  the  young  pro- 
glottids.  In  longitudinal  sections  certain  heavy  muscle  fibers  are  seen 
to  pass  from  the  neck  to  the  surface  of  the  head  near  the  tip,  others 
(the  greater  number)  pass  in  bundles  to  the  lower  and  inner  surfaces  of 
the  suckers.  A  vertical  muscle  cross  connecting  the  two  adjacent  suckers 
by  the  lower  margin  of  each  sucker  to  the  upper  margin  of  the  other  is 
also  distinguishable.  The  muscles  of  the  head  have  not  been  studied  in 
many  of  the  species.  The  writer  has  studied  them  in  Ophiotaenia  fila- 
roides,  Proteocephalus  amhloplitis,  and  P.  singularis  and  to  a  certain 
extent  in  P.  pinguis.  The  muscles  of  the  head  of  P.  singularis  are  de- 
scribed in  another  part  of  this  work  (vide  infra)  mth  drawings  illus- 
trating the  structures  found. /Riggenbach  (1896)  has  studied  the  muscu- 
lature of  the  head  of  P.  fossatus  and  Coralloiothrium  lobosum.  In  the 
latter  he  noted  certain  variations  from  the  type  found  in  other  Proteo- 
cephalids.  These  variations*  are  associated  with  the  type  of  head  found 
in  that  genus. 

The  genital  pore  is  situated  on  the  lateral  margin  of  the  proglottid, 
right  or  left.  Its  position  alternates  irregularly  from  one  side  to  the 
other.  For  each  species  the  genital  pore  has  a  fairly  constant  location 
on  the  margin.  In  some  species  it  is  near  the  middle,  and  in  other  cases 
it  is  posterior  to  the  middle,  in  others  anterior.  A  genital  papilla  or 
eminence  on  which  the  pore  is  situated  is  not  usually  present.  In  one 
species,  Ophiotaenia  grandis,  there  is  frequently  a  marked  pitting  or 
contraction  about  the  genital  pore  which  causes  the  latter  to  be  deeply 
set  back  from  the  straight  line  of  the  margin.  The  genital  pore  leads 
into  the  genital  atrium  into  which  typically  both  cirrus  and  vagina  open 
and  to  which  the  outer  end  of  the  cirrus-pouch  is  attached.  The  atrium 
is  to  be  considered  as  an  invagination  of  the  outer  body  wall  of  the 
proglottid. 


25]  PROTEOCEPHALIDAE  —  LA  RUE  25 

The  cirrus-pouch,  cirrus,  ductus  ejaculatorius,  vas  deferens,  vasa 
efferentia,  and  testes  comprise  the  male  reproductive  system. 

The  cirrus-pouch  is  a  more  or  less  cylindrical  or  ovoidal  muscular 
bag  which  contains  the  cirrus  and  ductus  ejaculatorius.  At  its  outer  end 
it  is  attached  to  the  wall  of  the  genital  atrium  by  means  of  some  of  its 
muscle  fibers.  At  the  other  end  strong  muscle  fibers  attach  it  to  the 
inner  dorsal  surface  of  the  muscle  sheath.  Some  muscles  are  also  con- 
tinued into  the  cirrus  itself.  The  form  and  size  of  the  cirrus-pouch  is 
reasonably  constant  for  each  species  if  proglottids  of  the  same  degree  of 
development  be  considered.  Hence  the  length  of  cirrus-pouch  relative 
to  the  breadth  of  the  proglottid  is  useful  as  a  diagnostic  character.  In 
some  instances  the  cirrus-pouch  is  somewhat  constricted  about  the  mid- 
dle. This  condition  seems  to  be  correlated  with  the  act  of  protrusion,  as 
the  writer  points  out  under  a  later  caption.  The  greatest  diameter  is 
usually  found  at  the  inner  end  of  the  pouch. 

The  more  or  less  muscular  cirrus,  a  tube  the  two  ends  of  which  vary 
somewhat  in  structure,  passes  through  the  length  of  the  cirrus-pouch. 
The  outer  portion  of  the  tube  is  called  the  cirrus  while  the  inner  and 
less  muscular  end  is  called  the  ductus  ejaculatorius.  Each  part  is  char- 
acterized by  certain  structures  tho  the  one  grades  over  into  the  other 
almost  imperceptibly.  The  cirrus  possesses  both  circular  and  longitudi- 
nal muscles.  The  circular  muscles  are  most  heavily  developed  near  the 
base  or  outer  end  of  the  cirrus  while  they  are  greatly  reduced  in  the  end 
where  it  passes  over  into  the  ductus  ejaculatorius.  Longitudinal  muscles 
are  weakly  developed.  At  the  outer  end  strands  of  muscle  pass  out  from 
the  cirrus  to  the  wall  of  the  cirrus-pouch.  There  are  very  few  if  any 
gland  cells  on  its  exterior.  Its  internal  surface  is  lined  by  a  continuation 
of  the  cuticula  of  the  atrium.  Toward  the  inner  end  of  the  cirrus  the 
cuticula  gradually  disappears  until  at  the  inner  end  it  is  difficult  to  dis- 
tinguish. The  muscles  also  become  weaker  and  the  outer  diameter  of 
the  tube  may  consequently  diminish  in  size  tho  in  some  species  this 
region  may  be  inflated.  "With  the  change  in  the  musculature  and  the 
thinning  of  the  cuticular  lining  prostate  gland  cells  appear.  These  char- 
acters mark  the  ductus  ejaculatorius,  namely,  the  thin  wall,  weak  mus- 
cles, little  or  no  cuticula,  and  the  presence  of  the  prostate  gland  cells. 
This  region  may  be  swollen  into  a  voluminous  tube  for  the  retention  of 
spermatozoa  and  hence  with  the  coils  of  vas  deferens  may  serve  as  a 
vesicula  seminalis.  There  is  no  specialized  vesicula.  The  cirrus  itself 
is  usually  quite  straight  and  is  only  rarely  thrown  into  coils.  The  ductus 
ejaculatorius  is  frequently  straight  tho  in  a  large  proportion  of  the  spe- 
cies it  is  bent  or  coiled  in  one  to  several  coils. 


26  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [26 

Since  the  outer  end  of  the  cirrus  tube  is  fastened  to  the  mouth  of 
the  cirrus-pouch  the  cirrus  is  everted  when  protrusion  takes  place.  The 
protruded  cirrus  is  then  of  double  thickness,  being  made  up  of  the  heavier 
walled  outer  tube  and  the  thin  walled  inner  tube.  In  certain  species  of 
Ophiotaenia,  viz.,  0.  marenzelleri  and  0.  trimeresuri,  one  to  three  or 
more  coils  of  ductus  ejaculatorius  are  forced  out  into  the  swollen  base  of 
the  cirrus.  This  could  only  occur  in  species  in  which  the  ductus  has  a 
considerable  length.  Species  having  a  short  straight  ductus  would  not 
show  this  character.  In  the  cases  above  cited  the  cirrus-pouch  is  not  evag- 
inated  as  Scjiwarz  (1908)  stated  was  true  of  0.  marenzelleri.  This  con- 
dition of  the  cirrus  and  ductus  is  more  fully  discussed  under  the  descrip- 
tions of  0.  marenzelleri  and  0.  trimeresuri.  (See  also  Figs.  106,  108, 
199.)  Such  a  pushing  out  of  coils  of  ductus  into  the  cirrus  is  not  known 
among  the  species  of  Proteocephalus,  Choanotaenia,  Crepidobothrium,  or 
Monticellia.  The  writer  has  recently  seen  it  in  some  proglottids  of  a 
species  of  Ophiotaenia  from  a  king  snake  from  Florida. 

In  certain  species,  notably  P.  fallax  (Fig.  57)  a  set  of  strong  circu- 
lar muscles  and  a  constricted  area  may  be  fou^d  about  the  middle  of 
the  cirrus-pouch.  The  inner  end  of  the  pouch  may  also  be  swollen.  The 
presence  of  these  muscles  together  with  the  constricted  area  which  is  not 
usually  seen  and  the  swollen  inner  end  suggest  a  possible  explanation 
of  the  process  of  cirrus  protrusion.  In  all  species  the  circular  muscles 
of  the  sheath  or  pouch  are  fairly  well  developed  while  the  longitudinal 
muscle  fibers  have  a  weaker  development.  A  contraction  of  the  circular 
muscles  and  perhaps  also  of  longitudinal  muscles  would  produce  a  rela- 
tively high  pressure  upon  the  contents  of  the  cirrus-pouch.  The  inner 
end  of  the  pouch  is  closed,  'the  outer  end  is  opened  broadly  and  at  the 
edges  of  this  broad  opening  the  cirrus  walls  are  attached.  It  is  evident 
therefore  that  the  only  escape  for  the  contents  which  are  under  pressure 
is  by  means  of  this  open  end  of  the  pouch  through  which  the  cirrus  is 
everted.  As  an  aid  in  the  beginning  of  the  act  of  protrusion,  relaxation 
of  the  muscles  of  the  cirrus  and  a  contraction  of  the  weak  muscles  which 
connect  the  cirrus  to  the  cirrus-pouch  may  play  a  part.  This  action 
once  initiated  the  hydraulic  pressure  put  upon  the  contents  of  the  cirrus- 
pouch  by  the  contraction  of  the  muscles  of  the  pouch  would  complete 
the  act  of  eversion.  In  certain  cases  where  the  cirrus  and  the  ductus 
ejaculatorius  form  a  straight  tube  through  the  cirrus-pouch  it  seems 
probable  that  a  shortening  of  the  pouch  is  also  necessary.  Such  a  short- 
ening has  not  actually  been  observed. 

The  retraction  of  the  cirrus  is  not  so  easily  explained.  It  seems  that 
strong  contractions  of  the  longitudinal  muscle  fibers  within  the  walls  of 
the  cirrus  and   ductus  ejaculatorius  would  be  of  great  assistance  in 


27]  PROTEOCEPHALIDAE  —  LA  RUE  27 

retraction.  In  many  species  the  muscle  fibers  connecting  the  cirrus  with 
the  walls  of  the  cirrus-pouch  seem  entirely  too  weak  to  assist  much  in 
this  process.  Since  the  inner  end  of  the  cirrus-pouch  is  attached  to  the 
body  wall  by  strong  muscles  their  contraction  would  lengthen  the  cirrus- 
pouch  and  this  would  exert  a  direct  traction  upon  the  ductus  ejaculato- 
rius.  Such  a  lengthening  of  the  pouch  would  tend  toward  the  produc- 
tion of  a  negative  pressure  which  might  draw  the  cirrus  back  into  the 
pouch.  In  some  species  large  numbers  of  cirri  have  been  seen  protruded, 
in  others  none  or  almost  none.  Protruded  and  unprotruded  cirri  have 
been  noted  among  both  mature  and  ripe  proglottids.  These  facts  stimu- 
late one  to  inquire  whether  the  cirri  are  normally  retracted  after  pro- 
trusion? Is  protrusion  of  the  cirrus  a  common  occurrence  in  the  act  of 
copulation  or  is  it  an  accident?  Had  those  unprotruded  cirri  in  the 
ripe  proglottids  ever  been  protruded?  These  are  questions  that  cannot 
be  answered  from  the  available  data  nor  has  the  author  seen  anything 
in  the  literature  on  these  points.  An  opportunity  is  suggested  here  for 
experimental  work. 

/^iggenbach  (1896)  considered  that  the  broad  cavity  of  the  cirrus 
might  be  called  a  vesicula  seminalis  and  that  this  cavity  functioned  as 
such  an  organ.  While  it  is  true  that  the  cavity  of  the  cirrus  and  ductus 
ejaculatorius  might  function  as  a  vesicula,  they  are  not  structurally  a 
vesicula.  When  the  sexual  organs  become  fully  mature  the  semen  is 
stored  until  ejaculated  in  the  coils  of  the  vas  deferens  which  become 
greatly  swollen.  Thus  the  coils  of  vas  deferens  function  as  a  vesicula 
seminalis  altho  they  are  not.  differentiated  into  such  an  organ.  JProperly 
speaking  there  is  no  vesicula  seminalis  in  the  Proteocephalidae.  The 
vasa  efferentia  and  vas  deferens  have  been  carefully  described  by  Rig- 
genbach  (1896)  and  La  Rue  (1909).  Altho  they  have  been  observed  in 
a  number  of  species  of  the  family  they  have  not  been  especially  re- 
described  in  the  descriptions  of  the  species  because  they  are  essentially 
the  same  in  all  species  of  the  group.  Their  general  relations  may  be  seen 
in  figures  90  and  180.  The  vasa  efferentia  are  thin  walled  small  tubes 
leading  out  from  the  covering  of  the  testes.  These  tubules  anastomose 
with  some  of  their  neighbors  and  finally  the  larger  vessels  unite  with  the 
inner  end  of  the  vas  deferens.  The  vasa  efferentia  can  sometimes  be 
studied  from  toto  preparations  of  proglottids  if  the  testes  were  discharg- 
ing semen  at  the  time  of  killing  the  cestode.  Carefully  made  frontal 
sections  stained  with  haematoxylin  sometimes  show  them  very  well.  The 
vas  deferens  is  a  thin  walled  tube  of  larger  diameter  than  the  vasa  effer- 
entia. It  begins  near  the  middle  of  the  proglottid  on  the  inner  dorsal 
surface  of  the  muscle  sheath.  Then  it  makes  a  few  or  numerous  coils 
which  extend  to  the  inner  end  of  the  cirrus-pouch  which  it  enters.    From 


28  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [28 

this  point  the  vas  deferens  is  continuous  with  the  ductus  ejaculatorius 
and  the  cirrus. 

Testes  are  numerous  small  ovoidal  or  spheroidal  bodies  which  lie 
toward  the  dorsal  surface  of  the  medullary  parenchyma  within  the  lon- 
gitudinal muscle  sheath.  They  occupy  as  a  rule  nearly  the  entire  space 
between  the  lateral  vitellaria  anterior  to  the  ovary.  They  leave  a  small 
free  area  in  the  region  of  the  coils  of  the  vas  deferens.  In  the  genus 
Proteocephalus  the  testes  form  one  or  two  layers  which  extend  from  one 
vitelline  field  to  the  other.  In  one  species,  P.  torulosus,  they  extend  past 
the  ovary  to  the  posterior  margin  of  the  proglottid.  In  the  genus  Coral- 
lobothrium  the  testes  form  a  single  broad  field  anterior  to  the  ovary.  In 
Ophiotaenia,  Crepidobothrium,  and  Acanthotaenia  they  form  two  lateral 
fields,  one  on  either  side  of  a  free  median  zone.  In  rare  cases  some  scat- 
tered testes  may  be  found  in  this  median  zone.  In  these  genera,  which 
have  been  but  little  studied  by  the  section  method,  the  testes  seem  to  form 
a  single  layer  in  depth.  The  number  of  testes  varies  widely  from  about 
25  in  some  to  about  400,  the  upper  limit  for  other  species.  For  each 
species  there  seems  to  be  a  characteristic  number  which  is  not  constant 
but  varies  within  certain  limits.  This  range  is  usually  not  great,  being 
5-10-15  for  cestodes  with  the  smaller  number  of  testes  and  very  much 
greater  for  those  cestodes  which  have  very  numerous  testes.  Crepido- 
hothrium  gerrardii  has  from  about  200  to  400  testes,  the  range  being 
about  200.  In  mature  proglottids  the  testes  are  most  plainly  seen.  As 
the  uteri  develop  and  the  eggs  fill  the  uterine  pouches  the  testes  are 
pushed  aside  by  the  swollen  uteri  or  they  shrink  and  are  lost  to  view 
between  the  walls  of  the  uterine  pouches. 

The  organs  which  comprise  the  female  reproductive  system  are 
vitellaria,  vitelline  ducts,  vagina  in  all  its  parts,  ovary,  oocapt,  oviduct, 
ootype,  shell  glands,  receptaculum  seminis,  uterine  passage  and  uterus. 
The  vitelline  follicles  are  small  spheroidal  bodies  which  are  arranged  in 
two  long  bands  which  extend  the  full  length  of  the  proglottid  near  the 
lateral  margin  of  the  latter.  The  follicles  usually  are  rather  compactly 
grouped  about  the  central  conducting  tubule.  Rarely  the  vitellaria  ex- 
tend past  the  posterior  edge  of  the  ovarian  lobes,  and  in  only  one  known 
case,  (Proteocephalus  perplexus),  do  they  follow  along  the  posterior 
margin  of  the  proglottid  (Figs.  54,  55).  The  vitellaria  are  inside  the 
longitudinal  muscle  sheath  and  very  near  the  lateral  nerve  trunks.  The 
central  conducting  tubules  near  the  posterior  end  of  the  proglottid  are 
known  as  the  paired  vitelline  ducts  which  lead  to  the  middle  of  the  pro- 
glottid where  they  unite  to  form  the  common  or  unpaired  vitelline  duct. 
This  may  dilate  to  form  a  vitelline  receptacle.  From  the  vitelline  recep- 
tacle the  unpaired  duct  passes  backward  to  join  the  oviduct  just  before 


29]  PROTEOCEPHALIDAE  —  LA  RUE  29 

the  latter  enters  the  ootype.  The  course  of  these  duets  as  described  by 
von  Linstow  (1891)  and  Kraemer  (1892)  was  incorrect,  ^^iggenbach 
(1896)  and  Bendict  (1900)  established  their  true  course.  The  vitelline 
ducts,  paired  and  unpaired,  are  made  up  of  thin  tissue  possessing  no 
glandular  elements. 

The  vagina  is  a  tube  of  varying  dimensions  and  character  leading 
from  the  genital  atrium  to  the  middle  of  the  proglottid  and  thence  pos- 
teriad  to  the  interovarial  space  where  it  unites  with  the  oviduct  to  form 
the  fertilization  canal,  which  is  a  part  of  the  oviduct.  The  vagina  and 
cirrus-pouch  alike  open  into  the  genital  atrium.  The  initial  part  of  the 
vagina  may  lie  anterior  or  posterior  to  the  cirrus-pouch  and  the  opening 
of  the  vagina  may  have  either  of  these  positions,  but  in  some  species 
examinedj^pecially  for  this  point  the  opening  was  dorsal  to  the  cirrus- 
pouch,  yin  Proteocephalus  the  vagina  is  usually  anterior  to  the  cirrus- 
pouch  altho  there  are  a  few  species  in  which  it  is  regularly  posterior. 
In  species  of  Ophiotaenia  it  is  as  frequently  posterior  as  anterior  to  the 
cirrus-pouch,  but  in  0.  filaroides  alone  it  has  never  been  observed  in  the 
posterior  position.  In  Acanthotaenia  and  Crepidobothrium  the  vagina 
may  be  either  anterior  or  posterior.  The  vagina  on  account  of  its  vary- 
ing structure  may  be  subdivided  into  four  parts.  These  are  (1)  the 
initial  part  reaching  from  the  genital  atrium  nearly  to  the  middle  of  the 
proglottid,  (2)  the  tube  extending  back  to  the  interovarial  space  where 
its  structure  changes  to  (3)  the  receptaculum,  and  then  (4)  the  lower 
vagina  extending  from  the  receptaculum  seminis  to  the  oviduct.  These 
parts  are  distinguished  by  position  and  by  structure.  The  initial  part 
has  an  opening  into  the  genital  atrium.  It  is  lined  usually  by  a  cuticula 
which  in  some  species  may  be  traced  almost  back  to  the  receptaculum. 
The  initial  part  is  frequently  somewhat  broadened  out  and  is  somewhat 
muscular.  Near  the  opening  to  the  atrium  the  muscles  are  usually  de- 
veloped into  a  sphincter  muscle,  the  sphincter  vaginae.  This  may  have 
a  strong  development  (Figs.  106,  183)  or  may  be  very  weak  (Figs.  47, 
50,  57).  The  modifications  of  the  sphincter  serve  as  distinguishing  char- 
acters. Noticeable  on  the  initial  part  of  the  vagina  particularly  but 
also  occurring  on  almost  the  entire  length  of  the  vagina  are  numerous 
gland  cells. 

The  longitudinal  tube  of  the  vagina  presents  some  different  appear- 
ances from  the  initial  part.  It  is  usually  a  straight  or  wavy  thin  walled 
tube  of  large  or  small  diameter  depending  on  the  species  and  the  stage 
of  development  of  the  proglottid.  In  some  species  as  the  proglottid 
becomes  mature  or  especially  after  coitus  this  part  of  the  tube  is  much 
dilated.  In  all  species  the  musculature  of  this  part  of  the  tube  is  very 
weak  or  lacking  entirely.    Here  also  the  cuticula  of  the  initial  part  of 


30  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [30 

the  vagina  may  disappear  or  is  very  thin  and  diflSeult  to  determine.  In 
Proteocephalus  macrocephalus  and  a  few  other  species  the  vagina  is 
ciliated  throughout  almost  its  entire  length.  Just  after  the  long  tube  of 
the  vagina  enters  the  interovarial  space  or  in  some  species  just  anterior 
to  the  mid-piece  of  the  ovary  the  vagina  broadens  out  slightly  and  this 
is  followed  by  a  sudden  diminution  in  diameter  and  a  change  in  the 
histological  structure  of  the  tube.  This  broadened  part  is  known  as  the 
receptaculum  seminis,  the  presence  of  which  Riggenbach  (1896)  denied. 
Since  it  is  somewhat  different  histologically  and  also  since  it  serves  the 
function  of  a  receptaculum  it  may  well  be  called  a  receptaculum.  The 
writer  has  already  presented  his  views  on  this  matter  in  his  description 
of  the  organ  in  Ophiotaenia  filaroides  (La  Rue  1909).  Since  that  time 
he  has  observed  the  structure  in  several  other  species  and  sees  no  reason 
for  changing  his  view.  From  the  receptaculum  seminis  to  its  junction 
with  the  oviduct  the  vagina  may  be  spoken  of  as  the  lower  vagina.  This 
part  is  characterized  by  its  small  diameter,  thick  muscular  walls  and 
rich  investment  of  gland  cells.  It  is  not  ciliated  in  any  part  of  its  course 
in  any  species  studied  by  the  writer.  The  lower  vagina  frequently  de- 
scribe a  long  loop  or  several  coils  within  the  interovarial  space. 

'''^The  ovary  as  in  other  TetraphyUideans  is  situated  in  the  posterior 
end  of  the  proglottid,  is  bilobed  and  the  lobes  are  connected  by  a  mid- 
piece.  The  lobes  are  thin,  alate  or  club-shaped,  branched  or  made  up  of 
several  slender  blind  pouches,  or  the  branches  may  anastomose.  Rarely 
or  never  is  the  lobe  a  solid  mass  but  it  seems  to  be  made  up  of  more  or 
less  anastomosing  branches  or  tubes.  The  whole  ovary  is  closely  in- 
vested with  a  thin  membrane.  The  organs  of  the  interovarial  region  are 
taken  up  as  nearly  as  possible  in  the  order  of  occurrence,  beginning  with 
the  oocapt.  The  general  relations  of  these  structures  are  shown  in  draw- 
ings which  have  been  reproduced  (Figs.  99, 104).  The  oocapt  is  a  funnel- 
shaped  muscular  organ  attached  to  the  mid-piece  of  the  ovary  and  pres- 
ent in  all  the  species  examined.  It  is  made  up  of  circular  and  longitudi- 
nal muscle  fibers  and  is  surrounded  by  scattered  gland  cells.  In  action 
it  is  a  gulping  organ  which  by  its  rhythmic  contractions  forces  the  ova 
down  the  oviduct.  The  oviduct  is  a  thick-walled  tube  made  up  of  epi- 
thelial cells  and  around  these  are  circular  and  longitudinal  muscle  fibers. 
Some  gland  cells  are  scattered  along  its  entire  length  and  it  is  lined  with 
numerous  long  cilia.  In  its  course  from  the  oocapt  it  describes  one  or 
more  loops  or  coils  and  then  it  receives  the  lower  vagina  which  here  pours 
the  sperms  upon  the  ova.  The  oviduct  from  this  point  on  should  be 
known  as  a  fertilization  pass^|)».  This  passage,  which  is  relatively  short, 
discharges  into  the  ootype.  However,  just  before  entering  into  the 
ootype  the  oviduct  receives  the  unpaired  vitelline  duct  which  pours  out 


31]  PROTEOCEPHALIDAE  —  LA  RUE  31 

the  product  of  the  vitelline  glands.  The  ootype  is  a  slightly  elongated 
muscular  organ  of  small  size.  Its  muscles  are  much  heavier  than  those 
of  the  oviduct  and  they  represent,  perhaps,  a  higher  development  of  the 
oviductal  muscles.  About  the  ootype  is  found  a  large  number  of  long 
slender  cells  which  in  the  aggregate  have  been  called  the  shell-gland. 
Each  cell  evidently  discharges  by  its  own  minute  duct  which  may  be 
traced  well  down  between  the  muscles  of  the  ootype.  In  the  ootype  the 
egg  is  formed.  The  ootype  discharges  directly  into  a  duct  called  the 
uterine  passage.  This  duct  leads  forward  directly  or  after  a  few  coils 
have  been  described  and  discharges  the  formed  eggs  into  the  median  stem 
of  the  uterus  at  a  point  which  is  usually  anterior  to  the  posterior  end  of 
the  uterus  and  on  the  dorsal  side  of  the  latter.  In  structure  the  uterine 
passage  differs  from  the  oviduct  in  the  lack  of  strong  musculature.  Some- 
times it  is  difficult  to  find  any  musculature  at  all.  There  are  scattered 
gland  cells  and  numerous  nuclei  of  cells  which  are  of  an  epithelial  nature 
or  which  belong  to  cells  of  parenchymatous  origin.  These  structures  of 
the  interovarial  space  have  been  worked  out  by  Monticelli  (1891),  von 
Linstow  (1891),  who  made  numerous  errors;  by  Kraemer  (1892),  who 
likewise  made  some  errors.  Riggenbach  (1896)  did  very  careful  work 
and  so  also  did  Benedict  (1900),  altho  the  latter  erred  in  denying  the 
existence  of  a  uterine  passage  for  his  species.  More  recently  the  writer 
(La  Rjle  1909)  has  worked  out  all  these  points  in  considerable  detail. 

j/n  all  the  species  of  the  family  the  uterus  develops  as  a  median 
longitudinal  tube  (Fig.  105)  lying  in  the  medullary  parenchyma  just 
within  the  longitudinal  muscle  sheath  and  toward  the  ventral  side  of  the 
proglottid,  while  the  testes  occupy  the  field  toward  the  dorsal  side  of  the 
proglottid.  This  long  tube  extends  almost  from  the  anterior  end  to  the 
mid-piece  of  the  ovary.  The  stem  of  the  uterus  is  first  seen  as  a  rod  of 
deeply  staining  cells  which  seem  to  arise  from  differentiating  parenchyma 
cells  in  young  proglottids.  The  rod  after  a  considerable  multiplication 
of  cells  becomes  hollow  throughout  its  entire  length.  By  the  method  of 
outpocketing  described  by  La  Rue  (1909)  the  uterus  sends  out  a  few 
or  numerous  branches  to  the  right  and  left.  These  branches  begin  as 
rods  of  cells  which  rapidly  multiply.  Then  a  cavity  opens  through  the 
length  of  the  rod  somewhat  as  the  lumen  appears  in  smaller  blood  ves- 
sels, during  the  development  of  higher  animals.  The  rod  continues  to 
increase  in  length  and  the  lumen  to  extend  as  the  rod  lengthens.  A  few 
of  the  outpocketings  are  directed  ventrally  and  these  eventually  pierce 
the  ventral  body  wall  (Fig.  89)  and  thus  provide  opportunity  for  the 
discharge  of  eggs.  The  lateral  pouches  are  formed  prior  to  the  dis- 
charge of  eggs  into  the  uterus  and  in  all  cases  observed  the  pouches  in- 
creased in  size  so  rapidly  that  there  was  no  crowding  of  eggs  until  the 


32  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [32 

uterus  had  almost  attained  its  complete  development.  Those  workers 
who  ascribe  the  production  of  these  pouches  to  the  crowding  of  the  eggs 
have  evidently  failed  to  read  the  evidence  to  be  seen  in  the  developing 
uterus.  Likewise  those  who  ascribe  the  splitting  of  the  proglottid  down 
the  ventral  side  to  the  pressure  of  contained  eggs  have  failed  to  note  the 
fact  that  the  ventral  pouches  perforate  the  ventral  side  in  one  or  more 
places  before  the  pressure  of  eggs  is  great  enough  to  split  the  proglottid. 

The  number  of  pouches  on  either  side  varies  greatly  but  is  reason- 
ably constant  for  each  species.  Proteocephalus  longicollis  with  its  three 
pouches  (Fig.  167)  on  either  side  has  the  smallest  number,  while  some 
of  the  Ophiotaeniae  (Fig,  101)  have  as  many  as  thirty  or  more.  When 
filled  with  eggs  the  pouches  in  many  species  fill  almost  the  entire  ventral 
field  of  the  proglottid,  while  in  some  species  of  Ophiotaenia  they  appar- 
ently take  up  but  little  more  than  half  the  width  of  the  proglottid.  The 
pouches  come  to  lie  very  close  together,  so  that  only  small  septa  separate 
them.  Testes  are  pushed  laterally  or  nearly  obliterated  by  the  pressure 
of  filled  pouches.  The  vitellaria  likewise  are  nearly  obliterated  in  many 
species  while  the  ovarian  lobes  usually  shrink  considerably  at  this  time. 
The  splitting  of  the  ventral  surface  and  the  discharge  of  the  eggs  occurs 
soon  after.  The  wall  of  the  uterus  in  its  early  stage  of  development 
appears  to  be  made  of  an  epithelium  the  cell  boundaries  of  which  are 
indistinct  or  invisible,  altho  the  nuclei  are  very  prominent  and  are 
closely  packed  together.  In  the  later  stages  of  the  uterus  the  nuclei  are 
farther  separated,  the  outlines  of  the  cells  cannot  be  seen  at  all  even 
tho  the  membrane  in  which  the  nuclei  lie  is  thick  and  apparently  tough. 
Riggenbach  (1896:92)  says,  ''Eine  dicke  Membran  bildet  die  Wand  des 
ganzen  Fruchtbehalters.  Nach  aussen  lagern  derselben  stets  viele  kleine 
Zellen  auf,  die  zwar  kein  eigentliches  Epithel  bilden,  wie  es  bei  andern 
Cestoden  oft  der  Fall  ist,  jedoch  als  Reste  eines  solchen  anzusehen  sind. ' ' 
In  this  view  Riggenbach  may  be  correct. 

The  eggs  usually  have  three  membranes  altho  in  a  few  species  the 
eggs  have  been  recorded  as  having  two  membranes.  In  these  cases  it  is 
possible  that  a  third  membrane  was  overlooked.  The  outer  membrane 
of  eggs  that  have  been  discharged  into  water  is  usually  much  larger 
than  the  others.  It  is  thin,  hyaline  and  spheroidal  in  form.  Eggs 
preserved  in  utero  and  later  examined  do  not  show  this  membrane  at  all 
prominently  and  in  some  cases  it  might  easily  be  considered  as  a  part 
of  the  second  membrane.  The  outermost  membrane  is  smooth  in  out- 
line. The  only  known  exception  is  found  in  the  eggs  of  Ophiotaenia 
nattereri  which  Schwarz  (1908)  described  as  having  peculiar  branched 
booklets  about  its  periphery.  His  drawing  of  this  egg  is  reproduced 
(Fig.  194).     The  middle  membrane  is  usually  thick  and  granular  and 


33]  PROTEOCEPHALIDAE  —  LA  RUE  33 

but  little  larger  than  the  embryo.  The  third  or  inner  membrane  is  a 
delicate  but  tough  and  clear  membrane  which  is  not  always  easy  to 
demonstrate.  It  is  closely  applied  to  the  embryo  and  might  readily  be 
overlooked. 

CHARACTERS   OP  DIAGNOSTIC  VALUE 

Benedict  (1900:339)  in  his  discussion  of  these  forms  called  at- 
tention to  the  small  dependence  that  can  be  placed  upon  external  meas- 
urements when  used  alone  as  a  means  of  determining  species.  While  in 
the  main  he  is  correct  in  his  contentions  it  is  certainly  a  fact  that  such 
measurements  when  used  in  conjunction  with  other  data  may  be  of 
great  v^e  and  these  should  always  form  a  part  of  the  species  descrip- 
tion.[/These  measurements  should  include  the  dimensions  of  the  head 
and  suckers,  the  length  of  the  strobila  and  its  breadth  at  intervals  or 
the  breadth  of  proglottids  at  certain  stages  of  development.  The  dimen- 
sions of  proglottids  are  always  of  service,  and  of  less  importance  are 
the  dimensions  of  the  neck.  Data  as  regards  the  host,  the  locality  and 
the  habitat  of  the  host  are  always  of  value.  The  older  investigators  made 
much  of  these  data  and  justly  so  for  they  were  unable  to  make  a  study 
of  the  internal  anatomy  of  cestodes.  At  the  present  time  such  data  do 
not  seem  to  be  as  highly  valued  as  they  should.  The  anatomy  and  finer 
structure  of  the  internal  organs  furnish  the  most  valuable  characters  for 
diagnostic  purposes.  The  relations  of  the  external  genitalia,  of  the 
male  and  female  reproductive  organs  to  each  other,  the  number  and  size 
and  distribution  of  the  testes,  character  of  the  coils  of  the  vas  deferens, 
size  and  character  of  the  cirrus-pouch,  cirrus,  ductus  ejaculatorius,  and 
the  protruded  cirrus  are  all  of  value  for  diagnosis.  So  also  are  the 
relations  of  the  various  parts  of  the  vagina,  the  vitellaria,  ovary,  oviducts 
and  all  of  the  small  oi^gaiis-of  the  tfrterovarial  space.  Of  especial  value 
are  data  in  regard  to  the  pouches  of  the  uterus  and  the  measurements 
of  the  egg  membranes  and  the  embryo.  The  presence  of  muscular 
elements  and  cilia  may  at  times  be  used  as  distinguishing  features. 
Drawings  showing  the  essentials  of  external  and  internal  anatomy  should 
always  form  a  part  of  a  report  on  these  animals. 


34  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [34 


KEY  TO  THE  BETTER  KNOWN  GENERA  AND  SPECIES  OF 
PROTEOCEPHALIDAE 

1(70)     Proteocephalids  with  heads  which  have  no  folds  or  lappets 

of  tissue  about  the  suckers 2 

2(47)     Testes  lie  in  a  single  broad  field  between  vitellaria;  para- 
sitic in  freshwater  fish. 

Genus  Proteocephalus  Weinland  1858 3 

3(24)     Functional  fifth  sucker  lacking,  i.  e.  absent  or  vestigial 4 

4(15)     Testes  number  100  or  more _ „ 5 

5(  8)     Cirrus-pouch  extends  j^-J^  across  breadth  of  proglottid „     6 

6(  7)     Uterine  pouches  4  to  8  on  either  side;  ductus  ejaculatorius 

nearly  straight Proteocephalus  agonis  (Barbieri  1909) 

7(  6)     Many  uterine  pouches  on  either  side;  ductus  ejaculatorius 
forming  several  coils. 

Proteocephalus  fossatus  (Riggenbach  1896) 
8(  5)     Cirrus-pouch  extends  less  than  ys  across  the  breadth  of  the 

proglottid  __ „ 9 

9(12)     Genital  pore  situated  at  middle  of  lateral  margin  of  proglot- 
tid   „.. „ „.._ 10 

10(11)  Uterine  pouches  3  to  4  on  either  side;  testes  in  2  layers;  cir- 
rus-pouch extending  %-%  across  the  breadth  of  the 
proglottid ;  suckers  0.18-0.2  mm.  in  diameter. 

Proteocephalus  torulosus  (Batsch  1786) 
11(10)     Uterine  pouches  7.to  14  on  either  side;  testes  in  1  layer,  cir- 
rus-pouch extending  Ye-yi  across  the  breadth  of  the 
proglottid;  suckers  0.095-0.16  mm.  in  diameter. 

Proteocephalus  macrocephalus  (Creplin  1825) 
12  (  9)     Genital  pore  anterior  to  middle  of  lateral  margin  of  pro- 
glottid   13 

13(14)  Uterine  pouches  20  to  25  on  either  side;  vagina  anterior  to 
cirrus-pouch;  135-155  testes. 

Proteocephalus  perplexus  La  Rue  1911 
14(13)     Uterine  pouches  10  to  12  on  either  side;  vagina  posterior  to 
cirrus-pouch;  about  200  testes. 

Proteocephalus  sulcatus  (Klaptocz  1906) 

15  (  4)     Testes  number  less  than  100 _ 16 

16(17)     Suckers  with  pointed  apex  and  shallow  cavity. 

Proteocephalus  singularis  La  Rue  1911 
17(16)     Suckers  round  or  oval  with  smooth  contour 18 


35]  PROTEOCEPHALIDAE  —  LA  RUE  35 

18(19)     Vaginal  sphincter  very  long  and    strong;    eestode    large, 

robust Proteocephalus  amhloplitis  (Leidy  1887) 

19(18)     Vaginal  sphincter  short  and  weak;  cestodes  small „ 20 

20(23)     Testes  in  2  partial  or  complete  layers „ „ 21 

21(22)  Cirrus-pouch  extends  ^-34  across  the  breadth  of  the  pro- 
glottid ;  ductus  ejaculatorius  forms  several  coils ;  75-90 

testes Proteocephalus  filicollis  (Rudolphi  1802) 

22(21)  Cirrus-pouch  extends  about  yi  across  the  breadth  of  the 
proglottid;  ductus  ejaculatorius  nearly  straight;  44-52 

testes Proteocephalus  esocis   (Schneider  1905) 

23(20)     Testes  in  one  layer Proteocephalus  neglectus  La  Rue  1911 

24 (  3)     Functional  (well  developed)  fifth  sucker  present 25 

25(26)     Suckers  pointed  at  posterior  margin. 

Proteocephalus  cy clops  (von  Linstow  1877)  sp.  inq. 

26(25)     Suckers  not  pointed  at  posterior  margin 27 

27(42)     Testes  number  about  70  or  less._._ 28 

28(35)  Cirrus-pouch  extends  less  than  %  across  breadth  of  pro- 
glottid   29 

29(32)     Embryos  0.03  mm.  in  diameter  or  greater 30 

30(31)     Uterine  pouches  6  to  8  on  either  side;  30-35  testes  situated 

in  1  layer Proteocephalus  fallax  La  Rue  1911 

31(30)     Uterine  pouches  7  to  14  on  either  side;  55-60  testes  situated 

in  2  partial  layers Proteocephalus  dubius  La  Rue  1911 

32(29)     Embryos  much  less  than  0.03  mm.  in  diameter. 33 

33(34)     Uterine  pouches  9  to  14  on  either  side ;  35-40  testes;  cestodes 

very  small _ Proteocephalus  exiguus  La  Rue  1911 

34(33)     Uterine  pouches  4  to  9  on  either  side;  50-60  testes;  cestodes 

of  medium  size Proteocephalus  percae  (Miiller  1780) 

35(28)  Cirrus-pouch  extends  less  than  %  across  breadth  of  pro- 
glottid   36 

36(39)     Testes  in  1  layer _ 37 

37(38)  Cirrus-pouch  extends  Yz-Ya  across  the  breadth  of  the  pro- 
glottid; 10-14  uterine  pouches  on  either  side;  embryos 
measure  0.016-0.018  mm.  in  diameter;  cestodes  slender. 

Proteocephalus  pinguis  La  Rue  1911 
38(37)     Cirrus-pouch  extends  Va-V7  across  the  breadth  of  the  pro- 
glottid ;  6-12  uterine  pouches  on  either  side ;  embryos 
measure  0.0212-0.018  mm.  in  diameter;  cestodes  short 

and  robust Proteocephalus  cernuae  (Gmelin  1790) 

39(36)     Testes  in  2  layers „ _ 40 

40(41)     Uterine  pouches  10  to  16  on  either  side;  cestodes  very  small 

and  weak Proteocephalus  pusillus  Ward  1910 


36 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


[36 


41(40)     Uterine  pouches  3  (or  4)  on  either  side;  cestodes  large  and 

robust I*roteocephalus  longicollis  (Zeder  1800) 

42(27)     Testes  number  more  than  70 „ „.. 43 

43(44)     Fifth  sucker  armed  with  minute  hooks  or  spines. 

Proteocephalus  osculatus  (Goeze  1782) 

44(43)     Fifth  sucker  unarmed 45 

45(46)  Vagina  anterior  to  cirrus-pouch;  genital  pore  posterior  to 
middle  of  proglottid;  suckers  0.25  mm.  in  diameter. 

Proteocephalus  skorikowi  (von  Linstow  1904) 
46(45)     Vagina  posterior  to  cirrus-pouch ;  genital  pore  about  middle 
of  proglottid ;  suckers  0.5  mm.  in  diameter. 

Proteocephalus  pentastoma  (Klaptocz  1906) 
47  (  2)     Testes  lie  in  two  lateral  fields  between  viteUaria;  parasitic 

in  amphibians,  aquatic  snakes  and  lizards __ _..  48 

48(49)  Cuticula  of  head  or  of  head  and  parts  of  strobila  covered 
with  minute  spines. 

Genus  Acanthotaenia  von  Linstow  1903 

49(48)     Cuticula  of  neck  free  from  spines  or  hooks 50 

50(69)     Suckers  rounded  or  oval  in  outline. 

Genus  Ophiotaenia  La  Rue  1911 
51(52)     Vagina  always  anterior  to  cirrus-pouch. 

Ophiotaenia  filaroides  La  Rue  1909 

52(51)     Vagina  either  anterior  or  posterior  to  the  cirrus-pouch..„ 53 

53(62)     Genital  pore  anterior  to  the  middle  of  margin  of  proglottid...  54 

54(57)     Uterine  pouches  20  or  less  than  20  on  either  side_ 55 

55(56)  Head  0.24-0.25  mm.  broad;  suckers  0.12-0.15  mm.  in  diame- 
ter; testes  number  80-100;  cirrus-pouch  extends  about 
Yi  across  the  breadth  of  proglottid. 

Ophiotaenia  nattereri  (Parona  1901) 
56(55)     Head  0.54  mm.  broad;  suckers  0.27-0.3  mm.  in  diameter; 
testes  number  100-120;  cirrus-pouch  extends  about  ^ 
across  the  breadth  of  proglottid. 

Ophiotaenia  racemosa  (Rudolphi  1819) 

57(54)     More  than  20  uterine  pouches  on  either  side 58 

58(61)     Testes  number  about  100-150 59 

59(60)  Testes  number  100-108,  situated  in  two  narrow  lateral  fields 
not  near  vitellaria ;  several  coils  of  ductus  ejaculatorius 
in  base  of  protruded  cirrus. 

Ophiotaenia  trimeresuri  (Parona  1898) 
60(61)     Testes  number  90-160,  situated  in  two  broad  lateral  fields 
extending  close  to  vitellaria,  no  coils  of  ductus  ejacula- 
torius in  base  of  protruded  cirrus. 

Ophiotaenia  Ibnnhergii  (Fuhrmann  1895) 


37] 


PROTEOCEPHALIDAE  —  LA  RUE 


37 


61(58)     Testes  number  150-215 Ophiotaenia  perspicua  La  Rue  1911 

62(53)     Genital  pore  at  or  near  middle  of  margin  of  proglottid 63 

63(66)     Heads  1.5  mm.  or  more  in  breadth 64 

64(65)     Muscular  wall  of  suckers  of  nearly  constant  thickness  about 
entire  circumference;  testes  number  150-240. 

Ophiotaenia  marenzelleri  (Barrois  1898) 
65(64)     Muscular  wall  of  suckers  much  thickened  on  anterior  mar- 
gin ;  testes  number  about  200. 

Ophiotaenia  punica  (Cholodkovski  1908) 

66(63)     Heads  less  than  1.5  mm.  broad 67 

67(68)     Testes  number  130-160;  ductus  ejaculatorius  forms  several 
coils;  24-35  uterine  pouches  on  either  side. 

Ophiotaenia  calmettei  (Barrois  1898). 
68(67)     Testes  number  200-250 ;  ductus  ejaculatorius  nearly  straight ; 
40-60  uterine  pouches  on  either  side. 

Ophiotaenia  grandis  La  Rue  1911 
69(50)     Suckers  with  point  of  lower  margin  re-entrant  into  cavity  of 

sucker Genus  Crepidohothrium  Monticelli  1899. 

The  only  know  species  of  this  genus  is 

Crepidohothrium  gerrardii  (Baird  1860). 

70(1)       Heads  with  folds  or  lappets  of  tissue  about  suckers _  71 

71(72)     A  single  fold  of  tissue  partially  covering  the  base  of  suck- 
ers; head  conical Genus  Choanoscolex  La  Rue  1911. 

The  only  known  species  is 

Choanoscolex  ahscisa  (Riggenbach  1896). 
72(71)     Many  folds  or  lappets  of  tissue  about  suckers;   anterior 
surface  of  head  flattened;  suckers  set  in  flattened  end 
of  head Genus  Corallohothrium  Fritsch  1886. 


DESCRIPTION  OF  SPECIES 

In  this  section  are  described  the  species  of  the  following  genera: 
Proteocephalus,  sensu  strictu,  Choanoscolex  La  Rue  1911,  and  Ophio- 
taenia La  Rue  1911,  of  the  family  Proteocephalidae.  The  species  of 
Monticellia  La  Rue  1911,  belonging  in  the  new  family  Monticellidae,  have 
also  been  described.  Other  species  which  have  been  mistakenly  placed 
in  the  genus  Proteocephalus  have  been  considered  and  where  possible 
their  rightful  position  pointed  out.  The  species  of  Corallohothrium  and 
of  Acanthotaenia  have  not  been  re-described. 


38 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


[38 


PROTEOCEPHALUS  FILICOLLIS  (Rudolphi) 

[Figs.  15,  58-60] 

1782:  Taenia  from  Gasterosteus 

1786 :  Taenia  from  Gasterosteus 

1790:  Taenia  gasterostei 

1800:  Alyselminthus  gasterostei 

1802:  Taenia  filicollis 

1803 :  Halysis  gasterostei 

1810 :  Taenia  filicollis 

1819 :  Taenia  filicollis 

1844 :  Taenia  filicollis 

1845 :  Taenia  filicollis 

1845:  Taenia  ambigua 

1850 :  Taenia  filicollis 

1850 :  Taenia  ambigua 

1858 :  Proteocephalus  ambigua 

1858 :  Proteocephalus  filicollis 

1872:  Taenia  ambigua 

1889 :  Taenia  filicollis 

1902 :  Taenia  filicollis 

1905 :  Ichthyotaenia  filicollis 

1909:  Ichthyotaenia  ambigua 

1911 :  Proteocephalus  filicollis 

Specific  Diagnosis:  Charactt 
Jjength  as  much  as  35  mm.  or  more.  Breadth  0.8  mm.  Proglottids  few, 
15-25.  Segmentation  indistinct.  Proglottids  attached  by  their  full 
width.  First  proglottids  broader  than  long,  0.20  mm.  broad  by  0.075- 
0.080  mm.  long.  Mature  proglottids  about  0.18  mm.  long  by  0.370  mm. 
broad.  Ripe  proglottids  0.48-0.66  mm.  long  by  0.425  mm.  broad.  Head 
small,  0.090-0.120  mm.  broad  or  perhaps  a  little  more,  not  always  well 
set  off  from  neck.  Suckers  0.042-0.055  mm.  in  diameter.  No  fifth 
sucker  present.    Neck  to  first  traces  of  segmentation  about  1.0  mm.  long. 

Genital  organs  as  in  genus.  Genital  sinus  marginal,  irregularly 
alternating,  situated  near  middle  of  segment  on  a  papilla-form  promi- 
nence. Cirrus-pouch  about  0.130  mm.  long  by  0.032-0.040  mm.  broad, 
reaching  about  y^-%  across  the  breadth  of  the  proglottid.  Cirrus-canal 
much  coiled  in  cirrus-pouch.  Vas  deferens  forming  mass  of  eccentric 
coils.  Testes  in  two  layers,  about  75-90  in  all.  Vagina  anterior  to 
cirrus-pouch.  Sphincter  vaginae  weakly  developed.  Receptaculum 
seminis  present.  ViteUaria  made  up  of  quite  compact  follicular  masses, 
in  lateral  fields.     Ovary  bilobed,  connected  by  an  arched  mid-piece. 


0.  F.  Miiller 

1782 :28-29 

Batsch 

1786 :241-242 

Gmelin 

1790 :3079 

Zeder 

1800 :255 

Rudolphi 

1802a  :114-115 

Zeder 

1803 :334 

Rudolphi 

1810:106 

Rudolphi 

1819 :148 

Bellingham 

1844:317 

Dujardin 

1845 :583 

Dujardin 

1845:583 

Diesing 

1850:512 

Diesing 

1850:512 

Weinland 

1858 :53 

"Weinland 

1858 :53 

Grimm 

1872 :243-246 

Lonnberg 

1889 :15 

Schneider 

1902:84,86-87 

Schneider 

1905 :21-24 

Liihe 

1909 :33 

La  Rue 

1911 :473, 474, 475 

of  the  genus. 

Verv  small  cestodes. 

39] 


PROTEOCEPHALIDAE  —  LA  RUE 


39 


Lobes  spheroidal,  thick.  Uterus  when  fully  developed  possessing  5-7-8 
lateral  pouches  on  either  side.  Eggs  provided  with  three  membranes. 
Outermost  membrane  0.050-0.075  mm.  in  diameter,  embryonal  covering 
0.032  mm.,  embryo  0.027  mm. 

Habitat :    In  intestine  of  host. 


Host 

Locality 

Collector 

Authority 

Gasterosteus  aculeatus* 

Miiller 

Muller  (1782:28-29) 

«                  « 

Greifswald 

Rudolphi 

Rudolphi  (1802a:  1 14- 1 15) 

«                  i< 

Berlin 

Olfers 

Diesing  (1850:512) 

«                  « 

Paris 

Dujardin 

Dujardin  (1845:583) 

a                        (1 

Ireland 

Bellinghain 

Bellingham  (1844:317) 

<i                         « 

Grimm 

Grimm  (1872:243-246) 

Gasterosteus  pungitius 

Rennes 

Dujardin 

Dujardin  (1845:583) 

«                   i< 

Halle 

Nitsch 

Diesing  (1850:512) 

<4                                          i< 

Greifswald 

Creplin 

Diesing  (1850:512) 

M                                      M 

Reval,  Esthonia 

(Russian  province) 

Schneider 

Schneider  (1905:21-24) 

«                                      « 

Porkhala,  Finland 

Levander 

Schneider  ( 1902 :84, 86, 87) 

"                                      " 

Upsala 

Lonnberg 

Lonnberg  (1889:15) 

0.  F.  Miiller  (1782:28-29)  stated  that  he  had  seen  cestodes  from  the 
intestine  of  Gasterosteus.  He,  however,  did  not  describe  the  species  nor 
did  he  propose  a  name  for  it.  One  of  his  drawings  of  this  cestode  in  its 
natural  size  presents  so  little  detail  that  it  is  of  little  value.  Miiller 's 
Taenia  gasterostei  which  he  found  in  the  body  cavity  of  Gasterosteus 
LS  none  other  than  Schistocephalus  solidus.  Batsch  (1786:224)  and 
Schrank  (1788:49)  followed  Miiller  in  using  this  name  to  designate 
the  cestode  found  in  the  body  cavity  of  the  stichling.  Batsch  (1786: 
241-242)  gave  but  a  few  notes  on  Miiller 's  Taenia  from  Gasterosteus 
and  he  referred  to  Miiller 's  (1782:28-29)  article  and  figures.  Gmelin 
(1790:3079)  used  the  name  Taenia  gasterostei  to  designate  the  cestode 
found  in  the  intestine  of  Gasterosteus  aculeatus  by  Miiller.  Gmelin 's 
diagnosis  reads:  ''TAENIA  GASTEROSTEI  81.— T.  tenuissima, 
Cauda  obtusa.  Miill.  Naturf.  18.  p.  28.  t.  3.  /.  6.  7.  Habitat  in  gasteros- 
tei aculeati  intestinis,  minima,  articulis  nudo  oculo  vix  discernendis." 
This  name  is  unavailable  because  of  Miiller 's  prior  use  of  it  for  the 
cestode  of  the  body  cavity.  Zeder  (1800:255)  in  his  Nachtrag  used  the 
name  Alyselminthus  gasterostei  to  designate  this  form.  His  description 
is  not  at  hand.  Rudolphi  (1802a  :114-115)  gave  this  species  the  name 
Taenia  filicollis.     His  specimens  came  from  Gasterosteus  aculeatus  at 

*Type  host 


40  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [40 

Greifswald.    Since  this  name  is  here  used  for  the  first  time  Rudolphi's 
description  is  quoted  in  full: 

"Taenia  filicollis:  capita  globoso  distincto;  collo  longissimo  filiformi; 
articulis  ovariisque  quadratis. 

Taenia  Gasterostei,  Syst.  Nat.  p.  3079,  n.  81. 

Alyselminthus  Gasterostei  Zeders  Nachtrag,  S.  255.  Taf.  3.  Fig.  1-4. 

"Zu  dem,  was  aus  Goezes  nachgelassenem  Manuscript  bey  Zedern  gesagt  ist, 
habe  ich  wenig  hinzuzusetzen.  Ich  habe  diesen  Wurm  nur  sehr  selten  in  Darmka- 
nal  des  Stichlings  gefunden,  und  noch  immer  durch  einen  ungliicklichen  Zufall 
ohne  den  Kopf.  Der  Hals  ist  lang,  diinn  und  ungegliedert ;  die  Glieder  des  Kor- 
pers  bis  auf  das  letzte,  welches  stumpf  abgerundet  ist,  viereckig.  Da  die  Eyer- 
stocke  so  sehr  ausgezeichnet  sind,  wundert  es  mich,  dass  er  ihrer  gar  nicht  er- 
wahnt,  so  wie  auch  die  Worte,  Glieder  hat  der  Wurm  eigentUch  nicht,  mir  unver- 
standlich  sind,  da  die  Glieder  sehr  deutlich  sind.  Die  Eyerstocke  sind  viereckig, 
und  zwar  mit  hervorstehenden  Ecken,  so  dass  sie  auf  den  vier  Seiten  einen  hellen 
Fleck  ubrig  lassen,  da  sie  selbst  weiss  und  undurchsichtig  sind.  Jedes  Glied  ist 
also  von  dem  andern  durch  einen  hellen  Fleck  abgesondert,  und  an  den  Seiten 
des  Wurms  lauft  eine  unterbrochene  helle  Linie,  wodurch  diess  Thierchen  ein 
sehr  schones  Ansehen  bekommt." 

Zeder  (1803:334)  gave  a  brief  diagnosis  of  this  species  which  he 
called  Halysis  gasterostei.  He  referred  to  the  writings  of  Miiller 
(1782),  Batsch  (1786),  Zeder  (1800),  Gmelin  (1790).  He  stated  that 
its  habitat  was  the  intestine  of  Gasterosteus  aculeatus.  Rudolphi  (1810: 
106)  gave  a  diagnosis,  description,  and  synonymy  of  this  species.  His 
synonymy  shows  very  clearly  that  he  himself  considered  his  species  to 
be  identical  with  the  forms  which  earlier  workers  had  described  as  Tae- 
nia gasterostei  (the  intestinal  parasite)  and  Alyselminthus  gasterostei. 
Rudolphi 's  diagnosis  and  description  are  here  quoted : 

"Taenia  filicollis  R.  Taenia:  capite  subgloboso  discrete,  collo  longissimo, 
filiformi,  articulis,  ovariisque  quadratis.  Miiller  im  Naturforscher  St.  18.  p.  28. 
Tab.  3,  fig.  6.  7.  Taenia  ex  intest.  Gasterostei.  Batsch  Bandw.  p.  241,  n.  14. 
Gmel.  Syst.  Nat.  p.  3079,  n.  81.  Taenia  gasterostei.  Goeze  apud  Zederum  in  huj. 
Nachtrag,  p.  255.  Tab.  3.  fig.  1-4.  Alyselminthus  Gasterostei.  Rudolphi  in  Wied. 
Arch.  III.  I,  p.  114.  Taenia  filicollis.  Zeder  Naturg.  p.  334,  n.  10.  Halysis 
Gasterostei. 

"Hab.  In  Gasterostei  aculeati  intestinis  Goezius  Augusto,  ipse  aliquoties  sed 
capite  orbam,  lunio,  reperimus. 

"Descr.     Vermes  duos  tresve  pollices  longi,  fere  lineam  lati,  candidissimi. 

"Caput  subglobosum,  discretum,  osculis  .orbicularibus,  majusculis,  binis  tarn 
superioribus,  quam  inferioribus.  Collum  filiforme,  longissimum.  Corpus  planum 
articulis  anticis  minoribus,  reliquis  subaequalibus,  quadratis,  ultimo  rotundato. 
Ovaria  quadrangularia,  angulis  acutis  productis,  opaca,  ut  articulorum  tantum 
partes  inter  angulos  sitae  pellucidae  sint.     Vermis    inde  aspectus   lepidus,   inter 


.41]  PROTEOCEPHALIDAE  —  LA  RUE  41 

quoslibet  enim  articulos  maculae  pellucidae,  et  simul  canalis  lateralis  pellucidi 
species  oboritur.     Foramina  non  visa. 

"Obs.  I.  Goezius,  qui  ovaria  non  vidit,  vermem  proprie  non  articulatum  esse 
in  posthumis  reliquit,  ipse  vefo  banc  speciem  aeque  ac  uUam  aliam  articulatam 
observavi.  A  Bothriocephalo  solido,  in  Gasterostei  abdomine  hospitante,  omnibus 
prorsus  notis  diversissima. 

"Obs.  2.  Vermem  in  aqua  fluviatili  biduum  vixisse,  in  puteali  enecari. 
Goezius  observavit." 

Rudolphi  (1819:148)  adds  nothing  to  his  former  data.  Bellingham 
(1844:317)  in  his  catalogue  of  Irish  Entozoa  reported  the  finding  of 
Taenia  filicollis  in  the  intestine  of  Gasterosteus  aculeatus.  His  observa- 
tions added  nothing  to  our  knowledge  of  the  structure  of  the  worm. 
Dujardin  (1845:583)  reported  that  he  had  found  Taenia  filicollis  in 
Gasterosteus  aculeatus  at  Paris  in  1838.  His  description,  a  part  of 
which  seems  to  have  been  derived  from  Rudolphi  (1810)  reads: 

"Long  de  50  a  8o  mm.,  large  de  2  mm.  environ ;  tete  presque  globuleuse, 
distincte  et  portee  par  un  cou  tres-long,  filiforme; — sans  trompe; — articles  presque 
Carres,  contenant  des  ovares  ( ?)  opaques  egalement  carres,  qui  laissent  entre  eux, 
et  sur  les  bords,  des  intervalles  demi-transparents." 

"Rudolphi  et  Goeze  I'ont  trouve  en  Allemagne,  dans  le  Gasterosteus  aculeatus. 
Je  I'ai  trouve  aussi  dans  ce  poisson,  a  Paris,  en  1838;  j'y  ai  vu  des  oeufs  a  double 
enveloppe,  dont  I'externe,  mucilagineuse,  est  longue  de  o  mm,  06  a  o  mm,  10; 
I'enveloppe  interne,  globuleuse,  est  large  de  o  mm,  036,  et  les  crochets  de  I'embryon 
sont  longs  de  o  mm,  012." 

Dujardin  (1845:583)  also  described  a  form  from  Gasterosteus  pun- 
gitius  (laevis)  which  he  named  Taenia  amhigua.  His  specimens  were 
collected  at  Rennes,  France.    His  description  reads: 

"Long  de  6  mm,  large  de  o  mm,  5  a  o  mm,  8,  forme  de  quinze  a  dix-sept  articles 
peu  distincts  et  de  forme  tres-variable ; — tete  petite,  large  de  o  mm,  17,  sans  trompe 
et  sans  crochets  tantot  retractee  tantot  saillante,  quelquefois  globuleuse  et  separee, 
par  un  cou  tres-etroit,  quelquefois  a  quatre  lobes  distincts,  correspondant  aux 
ventouses  larges  de  o  mm,  068  a  o  mm,  07 ; — cou  tres-contractile  et  dilatable,  traverse 
par  quatre  canaux  larges  de  o  mm,  009; — premiers  articles  males  avec  les  orifices 
genitaux  irregulierement  alternes,  et  les  penis  longs  de  o  mm,  16,  larges  de  o  mm,  032, 
rides  transversalement ; — derniers  articles  informes,  remplis  d'oeufs  globuleux  a 
double  enveloppe ; — enveloppe  externe  mucilagineuse,  longue  de  o  mm,  053  a  o  mm, 
058; — enveloppe  interne  longue  de  o  mm,  034;  embryon  de  o  mm,  026,  avec  des  cro- 
chets de  o  mm,  0095." 

"Je  I'ai  trouve  plusieurs  fois  a  Rennes  dans  I'intestin  du  Gasterosteus  laevis. 
Son  nom  specifique  exprime  sa  ressemblance  avec  les  scolex  et  les  caryophylles." 

If  Dujardin 's  description  of  his  T.  filicollis  and  his  T.  ambigua  be 
compared,  it  will  be  noted  that  in  the  measurements  of  the  second  egg 
membrane  there  is  very  good  agreement.    The  variation  in  the  size  of 


42  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [42 

the  outer  e^  membranes  in  Proteocephalus  is  large,  hence  but  little 
value  can  be  given  to  that  measurement.  Unfortunately  Dujardin  failed 
to  give  the  measurements  of  the  embryo  in  either  case,  so  no  comparison 
in  that  respect  is  possible.  Dujardin  records  a  difference  of  0.0025  mm. 
in  the  length  of  the  hooks  of  the  embryos  of  the  two  species.  That 
measurement,  however,  is  difficult  to  make  with  accuracy  and  little 
weight  should  be  given  it  as  a  means  of  differentiating  the  species.  It 
is  probable  that  Dujardin 's  specimens  from  Gasterosteus  aculeatus  and 
from  G.  pungitius  belonged  to  the  same  species.  Weinland  (1858:53) 
made  Taenia  ambigua  Dujardin  the  type  of  his  genus  Proteocephalus 
and  with  that  species  he  included  Taenia  filicollis  and  Taenia  dispar. 

Diesing  (1850:512)  recording  Taenia  filicollis  Rud.  added  nothing 
to  Rudolphi's  diagnosis  and  description  of  that  species.  He  gave  a 
complete  synonymy  up  to  his  time,  and  stated  that  this  species  had  been 
found  in  the  intestine  of  Gasterosteus  aculeatus  in  August  by  Goeze 
and  in  June  at  Greifswald  by  Rudolphi;  in  September  at  Berlin  by  de 
Olfers;  at  Paris  by  Dujardin;  in  Gasterosteus  pungitius  at  Halle  by 
Nitsch;  at  Greifswald  by  Creplin.  Concerning  Taenia  ambigua  Dies- 
ing (1850:512)  gave  nothing  new. 

Grimm  (1872:243-246)  found  a  form  in  Gasterosteus  aculeatus 
which  he  identified  as  Taenia  ambigua  Dujardin.  Grimm  published  no 
drawings  of  his  specimens.  The  important  parts  of  his  description  are 
here  quoted: 

"TAENIA  AMBIGUA  Duj. : — Im  Darme  des  bei  uns  so  haufig  vorkommenden 
Stichlings  (Gasterosteus  aculeatus),  von  denen  ich  in  vergangenen  Sommer  mehr 
als  loo  aufgeschnitten  habe,  fand  ich  unter  andern  6  Exemplare  einer  Bandwurm- 
gattung.  Sie  unterscheiden  sich  sehr  leicht  von  alien  ihren  Verwandten  und  sind 
augenscheinlich  zu  der  vori  Dujardin  aufgestellten  Species  Taenia  ambigua  zu 
rechnen,  obgleich  die  Bestimmung,  einer  kurzen  Beschreibung  des  ausseren  Habitus 
nach,  hochst  schwierig  ist  und  sehr  leicht  zu  einen  Irrthum  fiihren  kann. 

"Das  gn"osste  von  mir  aufgefundene  Exemplar  hatte  30  Mm  Lange  und  i  Mm 
Breite.  Die  Grossenverhaltnisse  der  einzelnen  Korpertheile  eines  11,5  Mm  langen 
Exemplars  sind  folgende;  die  Lange  des  Kopfchens — 0.13  Mm,  dessen  Breite — 
0,25  Mm,  dessen  Dicke — 0,15  Mm,  der  Durchmesser  der  Saugnapfe — 0,25  Mm,  die 
Lange  des  Raises — 1,1  Mm,  die  Lange  der  ersten  unreifen  Glieder  0,014  Mm; 
dieselben  verlangern  sich  allmalig,  indem  sie  0,02  Mm.,  0,03  Mm  u.  s.  w.  lang 
werden;  reifere  Glieder,  ungefahr  aus  der  Mitte  des  Wurms,  haben  eine  Lange 
von  0,20  M.,  und  das  vollkommen  reife  vorletzte  Glied  ist  0,43  Mm.  lang.  Die 
Dicke  des  Wurmkorpers  steigt  bis  zu  0,5  Mm. 

"Das  Kopfchen  des  Wurms  \x>n  der  Seite  betrachtet,  erscheint  etwas  ange- 
schwoUen ;  wenn  man  es  von  der  Flache  betrachtet,  so  sieht  man  an  den  in  Wein- 
geist  conservirten  Exemplaren  eine  Furche,  die  uber  den  Kopf,  an  dessen  freiem 
Ende,  von  der  einen  flachen  Seite  des  Wurms  zur  andern  verlauft.  Diese  Furche 
ist  aber  gewiss  keine  constante  Bildung,  wird  iber  dadurch  hervorgerufen,  dass 


43]  PROTEOCEPHALIDAE  —  LA  RUE  43 

der  Wurm,  uberhaupt  sehr  contractu,  sein  Kopfchen  ofters  einzieht,  namentlich 
wenn  er  in  Weingeist  gelegt  wird.  Einen  Riissel  besitzt  er  nicht.  Die  Saugnapfe, 
die  eine  runde  Form  haben,  sind  paarweise  auf  den  den  flachen  Seiten  des  Kor- 
pers  entsprechenden  Kopftheilen  angeordnet.  Die  Geschlechtsoffnungen  liegen  am 
Rande  der  Glieder,  eine  uber  der  andern.  Alle  Glieder,  die  mit  dem  Alter  all- 
mahlig  breiter  werden,  haben  eine  ziemlich  unregelmassige  Form,  besonders  aber 
ihre  Rander.  Die  Farbe  des  Wurms  ist  rein  weiss.  Indem  wir  noch  hinzusetzen, 
dass  das  letzte  died  an  seinem  hintern  Rande  bestiindig  etwas  aufgeschlitzt  ist, 
haben  wir  eine  moglichst  vollstandige  Darstellung  des  Aeussern  des  Bandwurms 
geliefert. 

"Dieser  Bandwurm  ist  hochst  zart  ....  — 

"Von  dem  weiblichen  Geschlechtsorgan  habe  ich  nur  die  Keimdruse  und  den 
Uterus  aufgefunden ;  die  Keimdruse  liegt  in  der  Mitte  des  Gliedes  und  hatte  in 
dem  untersuchten  Stadium  ein  spindelformiges  Aussehen ;  von  ihr  geht  der  Uterus 
aus,  der  aus  einem  Rohre  mit  feinen  faserigen  Wandungen  besteht,  das  mehrere 
Schlingen  bildet.  Der  Uterus  verlauft  in  die  Scheide,  die  sich  zum  Seitenrande 
des  Gliedes  begiebt,  wo  sie  nach  aussen  mit  einer  ziemlich  grossen  Oeffnung 
miindet. 

"Neben  dem  weiblichen  Geschlechtsporus  liegt  eine  andere  OeflFnung,  die  in 
das  mannliche  Geschlechtsorgan  fiihrt.  Das  letzte  besteht  aus  dem  kolbenformigen 
Cirrusbeutel  mit  dem  nach  innen  gewendeten  Cirrus,  der  an  seinem  Ende  etwas 
anschwillt.     Hinter  dem  Cirrusbeutel  liegt  die  doppelte  Samendrtise. 

"In  dem  letzten,  also  am  meisten  entwickelten  Gliede  waren  nur  Keime,  nicht 
aber  entwickelte  Eier  zu  sehen." 

Von  Linstow  (1878:231)  reported  as  hosts  for  Taenia  filicolUs  and 
Taenia  amhigua  only  those  which  had  been  reported  by  Diesing  (1850). 
Zsehokke  (1884:16-17)  reported  and  briefly  described  a  eestode  from 
Perca  fluviatilis,  Lake  Lucerne,  which  he  considered  to  be  Taenia  fili- 
collis  End.  His  specimens  probably  belonged  to  Proteocephalus  dubius 
La  Rue.  Leidy  ( 1886 :62-63 )  thought  that  he  had  found  Taenia  filicollis 
in  the  intestine  of  Amia  calva  from  North  Carolina.  His  specimens 
probably  belonged  to  the  species  for  which  La  Rue  (1911)  proposed  the 
name  Proteocephalus  perplexus  La  Rue.  Lonnberg  (1889:15)  reported 
the  finding  of  Taenia  filicollis  in  Gasterosteus  pungitius  at  Upsala.  He 
added  a  very  little  to  Rudolphi's  diagnosis. 

Kraemer  (1892)  reported  and  described  a  form  from  the  intestine 
of  Coregonus  fera,  Lake  Lucerne,  which  he  first  identified  as  Taenia 
filicollis  Rud.  In  the  same  paper  Kraemer  showed  that  his  Taenia  fili- 
collis and  his  Taenia  ocellata  were  identical.  Riggenbach  (1896)  and  Ben- 
edict (1900)  accepted  Kraemer 's  determination.  This  species  has  been 
shown  to  differ  from  the  Taenia  filicollis  of  Rudolphi  and  is  described 
under  the  name  Proteocephalus  fallax  La  Rue  1911,  where  a  full  dis- 
cussion of  the  position  of  Kraemer 's  species  may  be  found.    Benedict 


44  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [44 

(1900)  described  an  American  species  of  cestode  under  the  name  Pro- 
teocephalus  filicollis  (Rud.).  In  a  note  he  changed  the  name  to  Proteo- 
cephalus  ocellata  (Rud,).  He  accepted  Kraemer's  statement  that 
Rudolphi's  Taenia  ocellata  and  T.  filicollis  were  identical.  For  this  form 
which  proves  to  be  a  new  species  La  Rue  (1911)  proposed  the  name, 
Proteocephalus  exiguus  La  Rue.  Schneider  (1902:21-23)  reported  a 
parasite  from  Perca  fiuviatilis,  in  Finland,  as  Ichthyotaenia  filicollis 
(Rud.).  This  form  he  later  (1903:13  and  1905:15-17)  considered  to  be 
identical  with  0.  F.  Miiller's  Taenia  percae.  Schneider  (1902:84,  86- 
87)  reported  a  specimen  from  Gasterosteus  pungitius  as  Ichthyotaenia 
filicollis  (Rud.).  In  a  later  paper  (1905:21-24)  he  considered  this  form 
to  be  Ichthyotaenia  ambigua  Dujardin  and  here  he  gave  the  first  good 
description  of  this  species.  LUhe  (1909)  gave  a  diagnosis  of  this  species 
under  the  name  Ichthyotaenia  anihigua  Dujardin.  His  diagnosis  added 
nothing  to  the  data  given  by  Dujardin  (1845)  and  Schneider  (1905). 
La  Rue  (1911:473,  474,  475)  presented  some  of  the  conclusions  that 
are  given  in  the  present  paper  at  greater  length. 

The  name  Taenia  gasterostei  as  proposed  by  Gmelin  (1790)  for  this 
species  is  not  available  because  it  was  first  used  by  Miiller  (1782)  to 
designate  the  species  now  known  as  Schist ocephalus  solidus.  For  the 
same  reason  the  name  Alyselminthus  gasterostei  Zeder  (1800)  is  ren- 
dered invalid.  The  name  Taenia  filicollis  Rudolphi  (1802)  being  first 
used  to  indicate  this  species  remains  available.  The  generic  name  is 
Proteocephalus  as  indicated  by  "Weinland  (1858).  It  is  of  importance 
here  to  determine  the  systematic  position  of  the  specimens  reported  by 
Grimm  (1872)  and  by  Schneider  (1902  and  1905).  A  careful  compari- 
son of  their  descriptions  with' Dujardin 's  (1845)  description  of  Taenia 
amhigua  shows  that  they  evidently  belong  to  the  latter  species.  One 
notes  that  Grimm's  material  was  found  in  Gasterosteus  aculeatus  while 
Schneider's  specimen  came  from  G.  pungitius,  and  a  consideration  of 
these  data  at  once  precipitates  the  question  as  to  the  possibility  that 
T.  amhigua  Dujardin  and  T.  filicollis  Rudolphi  might  not  prove  to  be 
identical  and  hence  synonymous.  The  answer  to  this  question  must  be 
sought  in  the  study  of  the  collection  records  and  in  a  comparison  of  the 
structures  of  the  forms  involved. 

In  regard  to  habitat  one  may  present  certain  general  considera- 
tions that  should  have  weight.  The  identity  of  the  two  forms  can  not 
be  considered  proven  because  they  have  been  found  together  in  a  single 
host  or  separately  in  different  members  of  a  host  species,  for  numerous 
instances  could  be  cited  in  which  a  species  or  an  individual  of  that 
species  served  as  host  for  two  or  more  closely  related  species  of  cestode. 
Nevertheless,  it  is  true  that  a  knowledge  of  the  host  is  considered  to 


45]  PROTEOCEPHALIDAE  —  LA  RUB  45 

have  an  important  bearing  on  the  proof  of  an  identity.  Frequently  the 
knowledge  that  two  forms  have  the  same  species  for  a  host  furnishes 
the  first  intimation  that  they  may  prove  to  be  identical. 

One  may  also  make  the  statement  that  the  finding  of  specimens 
apparently  belonging  to  the  same  species  in  two  or  more  unre- 
lated host  species  may  give  an  early  intimation  that  these  specimens 
i>elong  to  different  species.  While  it  is  true  that  certain  cestodes  seem 
to  be  limited  in  their  habitat  to  a  single  host  species  there  is  a  sufficiently 
large  body  of  facts  to  warrant  the  belief  that  in  general  cestode  species 
are  not  so  limited.  It  is  of  common  acceptance  that  closely  allied  hosts 
having  a  wide  distribution  and  inhabiting  the  same  regions  and  espe- 
cially when  they  have  similar  feeding  habits  very  frequently  harbor 
the  same  species  of  parasites. 

Gasterosteus  aculeatus  and  G.  pungitius  are  closely  allied,  have 
wide  distribution,  inhabit  the  same  region,  and  have  similar  feeding 
habits.  Taenia  ambigua  has  been  reported  from  G.  aculeatus  by  Grimm 
and  from  G.  pungitius  by  Dujardin  and  Schneider.  Taenia  filicollis 
has  been  reported  from  G.  aculeatus  by  Rudolphi,  Goeze,  Diesing,  Du- 
jardin, and  Bellingham,  and  from  G.  pungitius  by  Diesing  (from  speci- 
mens collected  by  Nitsch  and  Creplin)  and  Lonnberg.  So  far  as  hosts 
are  concerned  there  is  evidence  to  believe  that  Taenia  filicollis  and  Tae- 
nia anibigua  are  identical  species.  Against  this  identity  is  the  fact  that 
Dujardin 's  description  of  T.  filicollis  indicates  a  larger  form  than  does 
his  description  of  T.  ambigua.  It  is  a  well-known  fact,  however,  that 
a  considerable  variation  in  length  and  breadth  may  exist  in  the  same 
species  of  cestode.  These  discrepancies  in  size  may  be  due  to  the  degree 
of  maturity  and  to  the  contraction  states  of  the  individual  worms.  It 
is  not  known  whether  Dujardin 's  specimens  of  T.  ambigua  had  at- 
tained their  full  development  nor  is  this  known  concerning  the  speci- 
mens described  by  Grimm  or  Schneider.  There  are  no  statements  to 
show  whether  Dujardin  measured  his  specimens  while  alive  or  after 
preservation.    This  would  make  a  real  difference. 

Attention  has  already  been  called  to  the  fact  that  in  regard  to  the 
egg  measurements  made  by  Dujardin  for  the  two  forms  in  question 
there  is  pretty  good  agreement.  There  is  not  sufficient  evidence  at  hand 
to  prove  positively  that  Taenia  ambigua  is  identical  with  Taenia  filicol- 
lis, yet  it  seems  that  the  evidence  for  the  identity  is  stronger  than  the 
evidence  against  it.  Therefore  in  this  work  the  writer  will  consider  that 
these  species  are  identical  and  since  Taenia  filicollis  has  the  right  of 
priority  that  name  should  be  retained.  It  is  to  be  hoped  that  this  dis- 
cussion, which  in  no  way  can  be  considered  to  settle  the  problem  of  the 
identity  of  these  species,  will  be  the  incitement  to  a  genuine  compara- 


46  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [46 

tive  study  of  the  cestodes  of  Gasterosteus  aculeatus  and  G.  pungitius. 
There  is  doubtless  a  large  amount  of  such  material  already  brought 
together  in  the  museums  and  private  collections  of  Europe  upon  which 
an  investigator  could  work. 

Schneider's  material  was  found  for  the  most  part  in  Gasterosteus 
pungitius  from  Obersee  near  Reval,  Esthonia  (a  province  in  Russia) 
in  the  months  of  May  and  June.  Never  more  than  four  cestodes  of  this 
species  were  found  in  a  single  host.  He  also  studied  a  single  specimen 
found  by  Dr.  K.  M.  Levander  in  Gasterosteus  pungitius.  This  speci- 
men was  reported  by  Schneider  (1902:84,  86-87)  as  Ichthyotaenia  fili- 
collis  (Rud.).  For  my  study  Dr.  H.  B.  Ward  very  kindly  secured  one 
of  Schneider's  specimens  of  this  species,  stained  in  carmine  and  mounted 
in  toto,  from  Dr.  K.  M.  Levander  in  Helsingfors.  It  bears  the  label, 
"Ichthyotaenia  ambigua,  int.  Gast.  pungitius  23/10  VI,  04."  The 
description  which  follows  is  based  on  Schneider's  (1905:21-24)  descrip- 
tion and  on  the  writer's  observations  on  this  one  mounted  specimen. 

Schneider's  longest  specimen  measured  as  much  as  35  mm.  long 
by  a  maximum  breadth  of  0.8  mm.  and  it  possessed  17  proglottids.  Of 
these  the  anterior  were  broader  than  long  and  they  were  united  by  their 
whole  width.  Proglottids  well  filled  with  eggs  are  longer  than  broad 
and  have  a  well  rounded  outline.  The  specimen  studied  by  the  writer 
has  about  25  proglottids.  Of  this  number  but  a  few  at  the  anterior  end 
are  young.  These  are  followed  by  a  large  number  of  nearly  mature 
and  mature  proglottids  and  these  in  turn  by  a  small  number  of  ripe 
proglottids.  The  strobilation  is  indistinct.  The  first  proglottids  meas- 
ure about  0.075-0.080  mm.  long  by  0.200  mm.  broad  while  the  mature 
proglottids  measure  about  0.180  mm,  long  by  0.370  mm.  broad  and  the 
ripe  ones  about  0.48-0.660  mm.  long  by  0.45  mm.  broad.  An  end  pro- 
glottid tapers  posteriorly  and  is  0.975  mm.  long  by  0.425  mm.  broad. 
Schneider  states  that  the  scolex  is  small,  0.100-0.120  mm.  broad,  and 
not  well  set  off  from  the  neck.  The  suckers  measure  0.055  mm.  in  diam- 
eter. No  fifth  sucker  is  present.  The  neck  is  relatively  broad  and  its 
length  is  about  one-fourth  that  of  the  body.  In  the  specimen  examined 
by  me  the  breadth  of  the  head  at  the  base  of  the  suckers  is  0.090  mm. 
It  is  very  short  and  not  well  set  off  from  the  rather  broad  neck.  Suck- 
ers are  about  0.042  mm.  in  diameter  over  all.  The  first  traces  of  seg- 
mentation occur  at  a  point  about  1.0  mm.  behind  the  head.  This  makes 
the  neck  equal  to  about  %  the  length  of  the  entire  body. 

It  is  stated  by  Schneider  that  the  excretory  vessels  are  quite  wide  ; 
that  the  two  ventral  vessels  which  communicate  in  each  segment  with 
the  exterior  through  a  submarginal  pore  measure  0.03  mm.  in  diameter ; 
and  that  the  dorsal  vessels  are  smaller,  their  diameter  being  0.0075  mm. 


47]  PROTEOCEPHALIDAE  —  LA  RUE  A7 

The  genital  pore  is  marginal,  irregularly  alternating,  and  situated  about 
the  middle  of  the  proglottid  length.  It  is  rather  prominently  situated 
on  a  little  eminence  which  is  most  evident  in  ripe  proglottids.  This 
eminence  may  be  caused  by  a  pushing  out  of  that  region  by  nearby 
organs. 

The  cirrus-pouch  measures  about  0.130  mm.  long  by  0.032-0.040 
mm.  broad.  Its  length  goes  into  the  proglottid  breadth  from  three  to 
four  times.  The  cirrus-canal,  as  Schneider  states,  is  greatly  coiled 
within  the  cirrus-pouch  and  it  broadens  out  at  iis  proximal  end  to  form 
a  vesicula  0.030  mm.  broad.  Coils  of  the  vas  deferens  form  a  mass 
which  lies  toward  the  porose  side  of  the  proglottid.  The  testes  in  the 
preparation  examined  by  the  writer  lie  in  two  layers  of  about  35-50 
testes  each.  Thus  the  total  number  of  testes  is  about  75-90  or  more. 
Schneider  found  that  the  testes  measure  0.055-0.062  mm.  in  sagittal  and 
0.035-0.045  mm.  in  frontal  diameter. 

The  vagina  lies  anterior  to  the  cirrus-pouch.  Schneider  found  a 
weak  sphincter  vaginae  near  the  vaginal  opening.  It  was  not  visible 
in  the  specimen  which, the  writer  studied.  The  vagina  passes  in  a  curve 
to  the  middle  of  the  segment  and  thence  posteriad  to  the  interovarial 
space.  Schneider  found  a  small  receptaculum  seminis  just  anterior  to 
the  ovary.  The  vitellaria  are  quite  compact  follicular  masses  in  the 
lateral  fields  of  the  segments.  Of  the  ovaries  Schneider  writes,  "The 
ovaries  appear  as  spherical  bodies  when  seen  from  the  surface.  In  the 
posterior  segments,  after  they  have  fiJ.led  the  uterus  with  eggs,  they 
wither  completely  or  the  remains  of  the  genital  organs,  pressed  together 
into  a  triangular  area,  are  recognizable  as  two  small  round  bodies  in  the 
posterior  region  of  the  segment.  I  hold  that  Grimm  (1872,  p.  246)  erro- 
neously considered  this  triangular  area  to  be  the  ovary  when  he  says: 
'the  ovary  lies  in  the  middle  of  the  segment  and  has  in  its  observed 
condition  a  spindle-like  appearance'  ".  The  ovary  is  bilobed,  the  some- 
what spheroidal  lobes  being  connected  by  a  short  arched  mid-piece.  In 
the  ripe  proglottids  observed  by  me  the  ovary  could  still  be  distin- 
guished. The  uterine  passage  was  found  by  Schneider  opening  into  the 
uterus  posterior  to  the  middle  of  the  proglottid  and  opposite  to  the  pre- 
formed opening  of  the  uterus  to  the  exterior.  When  immature  the 
uterus  is  a  median  tube  but  when  fully  developed  it  has  5-7-8  lateral 
pouches  on  either  side. 

As  observed  by  Schneider  the  outer  membrane  of  the  egg  has  a 
diameter  of  0.050-0.075  mm.  The  embryonal  covering  measures  0.032 
mm.  and  the  embrj^o  itself  0.027  mm.  These  measurements  of  the  egg 
agree  pretty  well  with  the  measurements  as  given  by  Dujardin  (1845). 
The  embryos  especially  agree  well  in  size,  and  it  may  be  pointed  out 
that  this  is  of  importance  in  seeking  to  prove  an  identity. 


48  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [48 

P.  filicollis  somewhat  resembles  P.  agonis  Barbieri  but  it  may  be 
differentiated  from  that  species  by  its  relatively  shorter  cirrus-pouch, 
by  a  different  position  of  the  mass  of  coils  of  vas  deferens,  by  its  larger 
testes  which  are  arranged  in  two  layers  while  the  testes  of  P.  agonis 
are  in  one  layer.  P,  agonis  has  no  genital  papilla  while  in  P.  filicollis 
there  is  a  simulation  of  one.  P.  filicollis  is  readily  distinguished  from 
P.  duhius  La  Rue  and  P.  fallax  La  Rue  by  its  smaller  size,  by  its  lack  of 
a  fifth  sucker,  by  its  more  numerous  testes,  shorter  cirrus-pouch  and 
smaller  proglottids.  The  embryos  of  P.  filicollis  are  smaller  than  those 
of  P.  fallax.  P.  filicollis  differs  from  P.  pusUlus  Ward  and  P.  exiguus 
La  Rue  in  not  having  a  fifth  sucker,  in  having  smaller  suckers,  a  smaller 
head,  more  testes,  and  fewer  uterine  pouches.  P.  filicollis  differs  greatly 
from  P.  percae  in  the  lack  of  a  fifth  sucker,  in  the  size  of  body,  head 
and  suckers,  in  the  proportions  of  the  proglottids,  in  the  size  of  the 
cirrus-pouch,  ovaries,  etc.  It  likewise  differs  from  P.  cernuae  in  prac- 
tically the  same  points  as  stated  for  P.  percae.  P.  filicollis  is  smaller 
than  P.  esocis.  It  has  a  shorter  cirrus-pouch  and  a  larger  number  of 
testes.  The  proportions  of  the  proglottids  are  different.  The  two  spe- 
cies are  alike  in  not  possessing  a  fifth  sucker. 


PROTEOCEPHALUS  ESOCIS   (Gui.  Schneider) 
[Figs.  16,  61-63] 
1905:    Ichthyotaenia  esocit  Gui.  Schneider,  1905:19-20 
1911:    Proteocephalus  esocis  La  Rue,  1911:475 

Specific  Diagnosis:  The  characters  of  the  genus.  Cestodes  small 
and  slender,  1-4.7  cm,  long,  maximum  breadth  0.7  mm.  Scolex  0.140 
mm.  broad,  0.105  mm.  thick.  No  fifth  sucker,  nor  vestige  of  it.  Suckers 
shallow,  weak,  small,  about  0.050  mm.  in  diameter,  directed  anteriad. 
Diameter  of  sucker  opening  0.026-0.032  mm.  Neck  2-5  mm.  long.  Sixty- 
six  proglottids  in  specimen  4.7  cm.  long,  youngest  proglottids  twice  as 
broad  as  long,  0.370  mm.  broad  by  0.170  mm.  long.  Middle  proglottids 
broader  than  long,  0.680  mm.  broad  by  0.390  mm.  long.  Last  proglottid 
longest,  0.650  mm.  long  by  0.460  mm.  broad  at  anterior  end.  Segmen- 
tation not  evident.  Transverse  intersegmental  furrows  shallow.  Sur- 
face smooth. 

Genital  organs  as  in  genus.  Genital  pore  not  on  genital  papilla. 
Pore  marginal,  irregularly  alternating,  about  middle  of  proglottid. 
Cirrus  sheath  relatively  long,  extending  nearly  to  middle  of  proglottid. 
Length  of  cirrus-pouch  0.230-0.250  mm.  Cirrus  sinuous.  Vas  deferens, 
a  mass  of  coils  in  middle  of  proglottid.    Testes,  30-38  visible  in  mature 


49]  PROTEOCEPHALIDAE  —  LA  RUE  49 

proglottids,  44-52  in  young  proglottids.  Testes  arranged  in  two  par- 
tial layers  between  vitellaria.  Diameter  of  testes,  0.050-0.090  mm.  Va- 
gina anterior  to  cirrus-pouch.  Near  vaginal  opening  a  small  sphincter 
vaginae.  Receptaculum  seminis  anterior  to  ovary.  Passages  in  inter- 
ovarial  space  as  in  genus.  Ovary  bilobed,  lobes  ovoidal,  vitellaria 
sparse,  follicles  small.  Uterus  not  observed.  Two  pairs  of  excretory 
vessels,  ventral  and  dorsal.    Dorsal  vessels  small,  ventral  vessels  much 

larger. 

Habitat:  In  intestine  of  Esox  lucius  L.  (type  host)  ;  Reval,  Es- 
thonia  (type  locality). 

This  species  was  described  and  delineated  by  Gjii.  Schneider  ( 1905 : 
19-20).  La  Rue  (1911:475)  included  it  in  a  list  of  species  belonging  to 
the  genus  Proteocephalus. 

This  species  from  Esox  lucius  taken  at  Reval  in  Esthonia,  a  prov- 
ince of  Russia,  Apr.  30,  1904,  is  briefly  described  by  Gui.  Schneider 
(1905).  Thanks  to  Dr.  H.  B.  Ward  the  writer  has  had  some  of  Schnei- 
der's preparations  and  alcoholics  for  examination.  The  paucity  of 
alcoholic  specimens  prevented  the  use  of  the  section  method.  The 
writer  has  therefore  been  compelled  to  be  content  with  a  study  of  toto 
preparations  and  specimens  cleared  in  glycerine.  The  description  fol- 
lowing is  based  largely  on  Schneider's  data  tho  wherever  possible  the 
writer  has  introduced  his  own  findings. 

The  worms  are  small  and  slender  1-4.7  cm.  long.  A  maximum 
breadth  of  about  0.7  mm.  was  observed.  No  worms  had  ripe  proglot- 
tids. The  diameter  of  the  scolex  is  about  0.140  mm.  The  four  shallow 
suckers  directed  partly  forward  are  weak,  small,  and  about  0.05  mm. 
in  diameter.  A  vestigial  fifth  sucker  is  lacking,  tho  perhaps  a  slight 
thickening  of  cells  in  the  anterior  part  of  the  head  is  to  be  considered 
as  the  vestige  of  the  shrunken  sucker  (Schneider).  My  measurements 
of  a  head  and  its  suckers  are :  breadth  0.140  mm.,  thickness  0.105  mm., 
diameter  of  suckers  0.053  mm.,  diameter  of  sucker  openings  0.026-0.032 
mm.  A  most  careful  search  of  Schneider's  toto  preparation  and  of  the 
specimen  cleared  in  glycerine  failed  to  reveal  a  trace  of  a  fifth  sucker 
or  vestige  of  it.  The  last  might  not  have  been  visible  under  these  con- 
ditions. A  functional  fifth  sucker  certainly  is  not  present.  The  suckers 
are  quite  prominent,  the  sucker  cavities  shallow.  The  slightly  de- 
pressed head  possesses  a  short  flattened  tip.  The  head  is  not  well  set 
off  from  the  neck.  Schneider  says  that  the  length  of  the  neck  is  about 
y3  the  length  of  the  body.    The  writer  finds  it  to  be  from  2.5-3-4.5  mm. 

Schneider's  largest  specimen,  4.7  cm.  long,  was  made  up  of  66 
proglottids.  The  youngest  proglottids  were  about  twice  as  broad  as 
long,  the  middle  ones  somewhat  broader  than  long,  and  the  posterior 


50  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [SO 

ones  quadrate  or  a  little  longer  than  broad.  The  intersegmental  fur- 
rows were  not  plain.  The  measurements  of  the  proglottids  in  a  glycer- 
ine specimen  examined  by  the  writer  were  as  follows:  First  proglot- 
tids 0.370  mm.  broad  by  0.170  mm.  long,  middle  and  mature  proglottids 
0.680  ram.  broad  by  0.390  mm.  long.  The  last  proglottid  in  the  strobila 
was  the  longest,  measuring  0.650  mm.  long  by  0.460  mm.  broad  at  ante- 
rior end.  The  segmentation  is  not  e\^dent.  The  proglottids  are  joined 
by  their  full  breadth  and  the  transverse  furrows  between  the  segments 
are  very  shallow.  No  longitudinal  furrows  are  present.  Drawings  of 
three  regions  (Figs.  61,  62,  63)  show  the  proportions  of  the  segments 
at  anterior,  post-middle,  and  posterior  region. 

The  genital  pore,  not  marked  by  a  papilla,  is  marginal,  irregularly 
alternating  in  the  strobila,  and  is  situated  about  the  middle  of  the  pro- 
glottid. The  cirrus-sheath  is  relatively  very  long  and  slender,  for  it 
extends  about  to  the  middle  of  the  proglottid.  Its  length  in  the  broad- 
est proglottids  is  about  0.230-0.250  mm.  Schneider  reports  that  its 
inner  end  lies  very  near  the  dorsal  muscle  layers.  This  is  also  the  case 
in  P.  percae,  P.  pitiguis  La  Rue,  P.  exiguus  La  Rue  and  a  number  of 
other  Proteocephalus  species.  The  cirrus  runs  through  the  cirrus-pouch 
in  a  sinuous  course  ^^^thout  the  formation  of  a  vesicula  seminalis.  The 
vas  deferens  forms  a  fairly  large  mass  of  coils  in  the  middle  of  the 
segment.  This  mass  however  is  much  smaller  than  in  P.  fallax  La  Rue 
or  in  P.  exiguus  La  Rue.  Schneider  found  about  30  testes  in  each  seg- 
ment, lying  in  a  single  layer,  about  8  in  a  transverse  row,  and  they 
measured  0.080-0.090  mm.  in  sagittal  by  0.040-0.050  mm.  in  frontal 
diameter.  It  has  been  found  difficult  to  count  the  testes  in  the  more 
mature  proglottids.  Thirty-eight  testes  are  shown  in  nearly  mature 
proglottids  (Fig.  62)  while  in  the  younger  proglottids  where  they  are 
smaller  and  more  easily  distinguished  the  writer  counted  44-48-52.  The 
testes  seemed  to  be  in  two  partial  layers  filling  the  area  between  the 
vitellaria.  The  larger  testes  measure  as  much  as  0.070-0.080  mm.  tho 
the  larger  number  measure  about  0.050  mm.  Schneider  found  the 
diameter  of  the  two  ventral  excretory-  vessels  to  be  about  0.015  mm. 
and  that  of  the  two  dorsal  about  0.0025  mm. 

The  vagina  (Fig.  62)  opens  anterior  to  the  cirrus-pouch.  Very 
near  the  opening,  0.030  mm.  from  it,  is  a  sphincter  muscle  0.015  mm. 
thick,  according  to  Schneider.  The  wTiter  was  unable  to  observe  this. 
From  its  opening  the  vagina  as  a  broad  tube  describes  a  long  smooth 
curve  to  the  middle  of  the  proglottid,  then  it  bends  posteriad  and 
passes  back  to  the  interovarial  space.  As  the  vagina  nears  the  middle 
of  the  segment  the  lumen  becomes  constricted.  Early  in  its  course  it 
crosses  the  cirrus-pouch  diagonally.    The  vagina  is  dorsal  to  the  uterus. 


51]  PROTEOCEPHALIDAE  —  LA  RUE  51 

Schneider  found  a  small  receptaculum  seminis  just  anterior  to  the 
ovary.  According  to  Schneider  the  passages  in  the  interovarial  space 
bear  the  relations  customary  to  Proteocephalids.  The  ovary  is  bilobed, 
the  lobes  being  like  two  elongated  balls  connected  by  a  slender  arched 
mid-piece.  The  vitellaria  are  lateral,  follicular,  sparse,  and  the  indi- 
vidual follicles  are  small.  The  uterus  was  not  observed  by  Schneider. 
It  is  not  well  developed  in  any  of  the  specimens  examined  by  the 
writer.  Schneider  saw  the  opening  of  the  uterine  passage  and  the  pre- 
formed uterus  opening  just  a  little  posterior  to  the  middle  of  the  pro- 
glottid.   No  eggs  were  seen.    No  proglottids  were  ripe. 

This  species  is  quite  closely  allied  to  the  P.  filicollis  (Rud)^ 
(amhiguus)  described  by  Gui.  Schneider  (1905).  However  it  is  differ- 
entiated from  P.  filicollis  by  its  much  longer  cirrus-pouch  and  by  its 
larger  testes.  P.  filicollis  (Schneider's  specimen)  shows  a  slight  genital 
prominence  upon  which  the  genital  pore  is  situated.  It  differs  very 
markedly  from  the  P.  pinguis  La  Rue  found  in  our  North  American 
Esox  lucius.  The  P.  pinguis  is  a  larger  worm  with  a  larger  head,  larger 
suckers,  a  functional  fifth  sucker,  a  relatively  shorter  cirrus-pouch,  and 
a  larger  number  of  testes.  P.  fallax  La  Rue,  P.  exiguus  La  Rue,  P. 
pusillus  Ward,  and  P.  percae  (0.  F.  Miiller)  are  readily  distinguished 
from  P.  esocis  by  means  of  the  fifth  sucker  and  many  other  character- 
istics which  are  best  shown  in  the  comparative  tables.  P.  macrocephalus 
(Creplin),  a  form  without  a  fifth  sucker,  is  much  larger,  has  more 
numerous  testes,  a  relatively  shorter  cirrus-pouch  and  in  other  ways 
also  it  differs  widely  from  P.  esocis  (Schneider). 

PROTEOCEPHALUS  AGONIS  (Barbieri) 
[Figs.  158,  174,  185] 

1909:    Ichthyotaenia  agonis  Barbieri,  1909:334-341 
1911:    Proteocephalus  agonis  La  Rue,  1911:475 

Specific  Diagnosis:  Characters  of  the  genus.  Cestodes  of  small 
size.  Strobila  3-4  cm.  long,  maximum  breadth  of  same  about  0.5  mm. 
Scolex  small,  0.168  mm.  in  diameter,  unarmed.  No  fifth  sucker.  Suck- 
ers circular,  symmetrically  arranged  on  head,  size?  Neck  quite  long, 
as  much  as  3  mm.,  breadth  0.140  mm.  Neck  not  well  differentiated 
from  strobila.  Proglottids  very  variable  in  form  and  number.  At 
times  as  many  as  50-70.  First  proglottids  broader  than  long.  Mature 
proglottids  longer  than  broad,  0.580-0.620  mm.  long  by  0.330-0.370  mm. 
broad.  Ripe  proglottids  filled  with  eggs,  nearly  quadrate,  length  and 
breadth  about  0.5  mm. 


52  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [52 

Genital  aperture  marginal,  irregularly  alternating,  situated  near 
middle  of  proglottid  length.  Male  organs  as  in  genus.  Testes  spher- 
ical, 0.031-0.038  mm.  in  diameter,  about  100  in  number.  Testes  packed 
in  area  between  vitellaria.  Coils  of  vas  deferens  in  middle  of  proglot- 
tid. Ductus  ejaculatorius  nearly  straight.  Cirrus  straight,  with  a 
thick  wall.  Cirrus-pouch  ovoidal,  broad  at  inner  end,  extending  to 
middle  of  proglottid.  Female  organs  as  in  genus.  Vagina  always  an- 
terior to  cirrus-pouch.  Uterus  in  ripe  proglottids  made  up  of  4-6-8 
lateral  pouches  on  either  side.    Diameter  of  eggs  0,037-0.038  mm. 

Habitat:  In  intestine  and  pyloric  coeca  of  Alosa  finta  var.  lacus- 
tris  Fa,  (type  host)  ;  Lake  Como,  Italy  (type  locality). 

The  material  was  collected  by  Prof.  Ciro  Barbieri  at  Bellagio  on 
Lake  Como  in  Italy  from  the  intestines  and  pyloric  coeca  of  Alosa  fiiita 
var.  lacustris  Fa.  He  found  them  here  in  great  numbers,  up  to  1200- 
1400  in  a  single  host. 

Barbieri  (1909)  described  this  species.  La  Rue  (1909:475)  in- 
cluded it  in  a  list  of  species  of  Proteocephalus.  Since  it  was  impossible 
to  secure  specimens  for  study  the  present  description  is  based  upon  Bar- 
bieri's  description. 

The  scolex  (Fig.  158)  is  very  small,  with  a  diameter  of  about  0.168 
mm.  It  is  unarmed  and  has  no  fifth  sucker.  The  four  suckers  are 
circular  in  outline  and  they  are  symmetrically  arranged  on  the  head. 
The  neck,  which  is  about  3  mm.  long  and  0.140  mm.  broad,  is  not  well 
differentiated  from  the  strobila.  The  proglottids  are  very  variable  in 
number  and  form.  One  well  developed  individual  with  ripe  eggs  in 
the  last  proglottid  had  50-70  proglottids.  The  first  proglottids  are 
broader  than  long.  Mature  proglottids  are  longer  than  broad,  0.580- 
0.620  mm.  long  by  0.330-0.370  mm.  broad.  Ripe  proglottids  filled  with 
eggs  are  nearly  quadrate,  length  and  breadth  being  about  0.5  mm.  The 
length  of  the  worm  is  variable.  Well  developed  individuals  measure 
30  mm.  long  while  a  single  individual  measured  40  mm. 

The  genital  aperture  is  marginal,  irregularly  alternating,  situated 
about  the  middle  of  the  length  of  the  proglottid,  and  marked  by  a 
shallow  depression.  The  testes  (Fig.  185)  fill  the  whole  area  between 
the  vitellaria  anterior  to  the  ovary.  They  are  nearly  spherical,  0.031- 
0.038  mm.  in  diameter.  Barbieri  figures  about  100  testes.  The  coils  of 
the  vas  deferens  form  a  thick  mass  which  occupies  the  middle  region 
of  the  proglottid.  Its  walls  are  thin  and  its  lumen  large.  The  ductus 
ejaculatorius  has  a  sinuous  course  in  the  inner  two-thirds  of  the  cirrus- 
pouch  but  apparently  it  forms  no  closely  twisted  coils  before  it  passes 
over  into  the  straight  heavy-walled  cirrus.     The  cirrus-pouch  extends 


53]  PROTEOCEPHALIDAE  —  LA  RUE  S3 

to  the  middle  of  the  proglottid.  It  is  ovoidal  with  its  broadest  part  at 
the  inner  end.    Its  wall  is  thick,  thickest  at  the  outer  end. 

The  ovary  is  bilobed,  the  lobes  being  oval  in  shape  with  smooth 
outlines.  The  vitellaria  are  lateral,  longitudinal,  follicular  glands 
which  lie  just  inside  of  the  lateral  ducts.  Oviduct,  ootype,  and  other 
organs  of  the  interovarial  space  are  as  in  other  species  of  the  genus. 
The  vagina  (Fig.  185)  always  lies  anterior  to  the  cirrus-pouch.  It 
widens  slightly  and  again  narrows  in  its  course  to  the  middle  of  the 
proglottid.  Near  the  inner  end  of  the  cirrus-pouch  it  bends  posteriad 
and  thence  takes  a  direct  course  to  the  interovarial  space  where  it  forms 
several  coils.  The  uterus  (Fig.  174)  containing  eggs  is  made  up  of 
4-6-8  lateral  pouches  which  in  fully  ripened  segments  fill  up  the  space 
between  the  vitellaria. 

The  excretory  system  is  of  the  usual  type.  Two  pairs  of  lateral 
excretory  ducts  take  their  origin  in  the  head  region.  They  empty  into 
a  small  bladder  situated  at  the  base  of  the  last  proglottid.  No  second- 
ary openings  were  observed  in  the  head  but  they  were  found  in  the 
posterior  region  of  each  proglottid. 

The  eggs  have  a  diameter  of  0.037-0.038  mm.  Barbieri  failed  to 
state  whether  this  measurement  included  only  the  embryo  or  embryo 
with  its  membranes.  Larval  stages  were  found  in  Bythrotrephes  and 
Leptodora. 

This  species,  as  will  be  seen  from  its  position  in  the  tables  at  the 
end  of  this  section,  is  most  closely  related  to  P.  esocis  (Schneider).  It 
is  differentiated  from  that  species  by  its  different  proportions,  by  its 
much  smaller  and  more  numerous  testes,  by  the  position  of  the  coils  of 
the  vas  deferens  and  by  the  relation  of  the  vagina  to  the  cirrus-pouch. 
P.  agonis  while  resembling  P.  filicollis  in  size  differs  from  that  species 
in  having  a  longer  cirrus-pouch,  in  the  excentric  position  of  the  coils  of 
vas  deferens,  and  in  having  smaller  and  more  numerous  testes  which 
lie  in  one  layer.  P.  agonis  is  readily  distinguished  from  P.  fallax,  P. 
duhius,  P.  pusUlus,  and  P.  exiguus  by  its  lack  of  a  fifth  sucker,  by  its 
more  numerous  and  smaller  testes,  and  by  its  less  numerous  uterine 
pouches. 


54 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


[54 


1900: 
1911; 


PROTEOCEPHALUS  EXIGUUS  La  Rue 

[Figs.  14,  50-52,  118,  135-138] 

Proteocephalus  filicolUs  Benedict,  1900:355-365 
Proteocephalus  exiguus  La  Rue,  1911 :477-478 


Specific  Diagnosis:  Characters  of  genus.  Strobila  short  and 
slender.  Length  9-15-25-38  mm.  Maximum  breadth  0.425-0.646-0.8 
mm.  Segmentation  not  evident.  Intersegmental  furrows  shallow. 
Neck  2-4-7-10 (?)  mm.  long  by  0.10-0.20  mm.  broad.  First  proglottids 
longer  than  broad  or  nearly  quadrate.  Mature  and  ripe  proglottids 
longer  than  broad.  Ripe  proglottids  considerably  larger  than  mature 
ones,  0.680-1.190  mm.  long  by  0.460-0.595  mm.  broad.  End-proglottid 
elongated,  pointed  posteriorly,  0.714-1.50  mm.  long  by  0.40-0.646  mm. 
broad.  Head  somewhat  globular,  flattened  dorsoventrally,  bearing  four 
suckers  on  its  broadest  part.  Breadth  of  head  0.120-0.160-0.170  mm. 
Suckers  0.058  mm.  broad  by  0.069-0.085  mm.  long.  Opening  of  suckers 
about  0.040  mm.  in  diameter.  Fifth  sucker  muscular,  functional,  0.037- 
0.048  mm.  in  diameter.  Common  genital  sinus  alternating  irregularly, 
situated  near  middle  of  lateral  margin  of  proglottid.  Testes  35-54  in 
number,  in  one  layer,  between  vitellaria,  anterior  to  ovary.  Diameter 
of  testes  0.04-0.05  mm.  Vas  deferens  forming  mass  of  coils  in  mid-field. 
Cirrus-pouch  0.289-0.340  mm.  long,  reaching  to  the  middle  of  segment. 
Cirrus  straight,  0.10  mm.  long  when  protruded.  Vagina  anterior  to 
cirrus-pouch,  crossing  cirrus-pouch  near  middle.  Sphincter  vaginae 
weak,  0.008  mm.  thick.  Lobes  of  ovary  retort-shaped  with  smooth  out- 
lines. Vitelline  follicles  small,  not  compact.  Uterus  when  fully  devel- 
oped with  9-14  lateral  pouches  on  either  side.  Embryos  about  0.019- 
0.021  mm.  in  diameter,  second  egg  membrane  0.036-0.046  mm.,  outer 
membrane  0.038-0.06  mm. 

Habitat:  In  stomach  (probably  by  post-mortem  wandering)  and 
intestine  of  host. 


Host 

Locality 

Collector 
H.  B.  Ward 

H.  B.  Ward 
H.  B.  Ward 

Authority 

Coregonus  nigripinnis 

C.  prognathus 
C.  artedi 

Lake  Michigan 
near  Charlevoix 
As  above 
As  above 

La  Rue 

La  Rue 
La  Rue 

Type :  Alcoholics  No.  Ch.  7c,  Ch.  2c,  and  an  unnumbered  bottle  in 
Doctor  H.  B,  Ward's  collection,  from  Coregonus  nigripinnis,  C.  progna- 
ihus,  and  C.  artedi.    Type  locality,  Lake  Michigan,  near  Charlevoix. 


55]  PROTEOCEPHALIDAE  —  LA  RUE  55 

Benedict  (1900:355-365)  described  some  cestodes  which  had  been 
collected  by  Dr.  H.  B.  Ward  while  engaged  in  a  biological  investigation 
of  Lake  Michigan  under  the  auspices  of  the  Michigan  Fish  Commission 
during  the  summer  of  1894.  The  hosts  were  Coregonus  nigripinnis,  C. 
prognathus,  and  C.  artedi.  Benedict  who  had  for  comparison  some 
specimens  identified  by  Zschokke  as  Taenia  ocellata  Rud.  (probably  P. 
fallax  La  Rue)  thought  his  material  and  Zschokke 's  specifically  iden- 
tical. He,  however,  called  attention  to  the  fact  that  Zschokke 's  speci- 
mens had  been  nearly  dried  out  in  transit  and  consequently  were  almost 
ruined  for  study.  Accepting  Kraemer's  statement  that  Taenia  filicollis 
Rud.  and  T.  ocellata  Rud.  were  identical  he  used  the  name  Proteocepha- 
lus  filicollis  (Rud.)  to  designate  his  form.  Later  he  discovered  that  the 
name  Taenia  ocellata  Rud.  had  page  precedence  in  the  original  paper 
by  Rudolphi  (1803)  and  in  a  foot-note  he  called  attention  to  the  fact 
that  the  name  Proteocephalus  ocellata  (Rud.),  mis-spelling  for  ocella- 
tus,  should  be  used  to  designate  the  form.  La  Rue  (1911:477-478) 
distinguished  this  species  from  P.  filicollis  and  described  it  as  a  new 
species  P.  exiguus. 

Careful  examination  of  prepared  specimens  from  some  of  the  same 
lots  used  by  Benedict  in  his  research  and  a  study  of  one  of  his  slides 
together  with  a  comparison  of  these  with  specimens  which  Zschokke  had 
sent  to  Dr.  H.  B.  Ward  under  the  name  of  Taenia  ocellata — (now  No. 
09.9  in  Professor  Ward's  collection) — has  convinced  the  writer  that  Bene- 
dict's species  is  a  new  one.  La  Rue  (1911)  proposed  for  it  the  name 
Proteocephalus  exiguus  La  Rue  by  reason  of  the  small  size  of  the  indi- 
viduals. Benedict  worked  out  the  morphology  of  the  form  quite  cor- 
rectly and  probably  would  have  given  due  weight  to  the  difference  be- 
tween his  own  and  Zschokke 's  specimens  had  the  latter  been  in  good 
condition. 

This  species  is  based  upon  the  work  of  Benedict  (1900:355-365) 
and  upon  a  further  examination  of,  and  collection  of,  data  from  certain 
lots  of  alcoholic  material  in  Dr.  H.  B.  Ward's  collection  from  which 
Benedict  secured  his  specimens  for  study.  These  bottles  bear  the  labels 
"Long-jaw,  stomach.  Charlevoix  Jy.  16,  '94".  "Ch.  7  c,  Cestodes,  Etc. 
Stomach  Black-fin  Charlevoix,  Jy.  16,  '94."  "Ch.  3  c,  cestodes.  Core- 
gonus Artedi  Jy.  12,  '94".  Slides  have  been  prepared  from  the  first 
two  lots. 

The  specimens  which  were  examined  carefully  by  the  writer  ranged 
in  length  from  9-10-15  mm.  Benedict  found  the  length  to  vary  from 
12-16-25  and  one  specimen  only  attained  the  length  of  38  mm.  This 
one  case  seemed  to  have  been  due  to  an  extreme  attenuation  for  the  neck 
was  7  mm.  long.    This  attenuation  may  have  been  caused  by  the  method 


56  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [56 

of  killing.  The  maximum  breadth  found  by  the  writer  varied  from 
0.425-0.595-0.646  mm.  while  Benedict  records  a  maximum  of  0.8  mm. 
The  globular  head  according  to  Benedict  is  0.120  mm.  broad.  The 
writer  found  it  to  be  0.148-0.160-0.170  mm.  broad.  The  suckers  placed 
just  above  the  broadest  zone  are  directed  slightly  anteriad.  They  are 
0.069-0.085  mm.  long  by  0.058  mm.  broad  with  an  opening  of  0.042-0.048 
mm.  by  0.027-0.032  mm.  Benedict  states  that  the  sucker  is  0.040  mm. 
in  diameter,  but  it  seems  that  he  must  mean  the  diameter  of  the  opening 
and  not  the  diameter  over  the  whole  sucker.  A  fifth  sucker  is  present, 
0.037-0.048  mm.  in  diameter.  This  from  toto  preparations  is  appar- 
ently a  true  sucker.  Benedict  says  it  has  the  **same  structure  and 
musculature  that  the  four  large  suckers  possess."  His  drawing  of  the 
head  is  reproduced  (Fig.  138). 

The  neck  measures  2-4  mm.  in  length  by  0.110-0.120  mm.  broad. 
Benedict  found  the  neck  to  be  5-10  mm.  long  by  0.10-0.20  mm.  broad. 
The  neck  passes  over  into  the  first  proglottids  almost  imperceptibly. 
The  first  proglottids  are  longer  than  broad  or  nearly  quadrate.  These 
very  immature  proglottids  are  few  in  number.  The  anlage  of  the 
genital  organs  appear  very  early  and  in  proglottids  a  little  further  back 
mature  sexual  organs  appear.  Mature  proglottids  are  longer  than  broad 
and  are  few  in  number.  Ripe  proglottids  are  longer  than  broad.  Three 
of  these  measured  0.935  by  0.595  mm.,  0.680  by  0.560  mm.,  1.190  by 
0.460  mm.  Ripe  proglottids  are  considerably  larger  than  the  mature. 
The  increase  in  length,  breadth  and  thickness  is  rapid.  The  longest 
proglottid  in  the  strobila  is  usually  the  last  one  and  sometimes  also  it 
is  the  broadest.  Three  terminal  proglottids  measured  1.190  by  0.646 
mm.,  0.714  by  0.51  mm.,  1.50  by  0.40  mm.,  the  length  being  stated  first. 
The  end  proglottid  is  variable  in  form  and  functional.  It  usually 
tapers  to  a  blunt  point  at  its  posterior  end  where  the  excretory  pore  is 
situated.  The  segmentation  is  not  evident.  The  angles  at  the  ends  of 
the  proglottids  are  scarcely  marked  and  the  intersegmental  furrows  are 
shallow.  The  total  number  of  proglottids  in  a  strobila  is  small,  10-12- 
14-15-20.    Benedict  found  one  specimen  with  40  segments. 

Benedict's  findings  on  points  of  histology  are  probably  correct  and 
have  not  been  checked  over.  The  common  genital  sinus  is  situated 
near  the  middle  of  the  margin  of  the  proglottid,  usually  slightly  ante- 
rior thereto.    It  alternates  irregularly. 

The  testes  (Fig.  50)  are  37-50-54  in  number,  35-50  according 
to  Benedict.  They  measure  0.040-0.045-0.050  mm.  in  diameter.  The 
testes  lie  in  a  single  layer  between  the  vitellaria  and  anterior  to  the 
ovarj'.  In  toto  mounts  the  writer  could  not  make  out  with  certainty  the 
coils  of  the  vas  deferens  yet  they  seemed  to  lie  in  the  middle  of  the 


57]  PROTEOCEPHALIDAE  —  LA  RUE  57 

proglottid  near  the  dorsal  wall.  Benedict  figures  the  vas  deferens  as 
forming  a  coil  extending  beyond  the  middle  of  the  segment.  The 
cirrus-pouch  (Figs.  50,  51,  52)  is  elongated,  slightly  broader  near  its 
inner  extremity  and  slightly  constricted  near  the  middle.  It  reaches 
the  middle  of  the  segment  or  even  beyond  that  point  in  mature  pro- 
glottids.  Its  length  increases  with  the  maturity  of  the  segment.  The 
following  measurements  of  its  length  were  taken  from  successive  pro- 
glottids,  0.289,  0.296,  0.313,  0.330,  0.341  mm.  the  last  being  the  maxi- 
mum length  found.  The  unprotruded  cirrus  is  straight.  The  writer 
has  not  seen  it  evaginated.  According  to  Benedict  the  evaginated  cirrus 
extends  0.10  mm.  from  the  male  opening.  The  ductus  ejaculatorius  iii 
straight. 

The  vagina  (Figs.  50,  51,  52)  opens  into  the  common  genital  sinus 
and  it  always  lies  anterior  to  the  cirrus-pouch.  In  a  single  proglottid 
the  vagina  was  found  posterior  to  the  cirrus-pouch.  This  is  probably 
to  be  considered  as  an  abnormality  or  a  very  rare  variation.  A  weak 
sphincter  vaginae  occurs  a  short  distance  from  the  vaginal  opening. 
This  sphincter  is  0.008  mm.  thick  and  is  circular  in  cross  section  (Bene- 
dict). As  Benedict  has  described  it,  the  vagina  extends  straight  in  for 
a  short  distance  then  it  "bends  backward  and  inward  passing  under 
the  middle  of  the  cirrus-sac".  The  ovary  very  much  resembles  that  of 
P.fallax  (Kraemer's  Taenia  ocellata)  in  shape.  The  lobes  are  thick  and 
solid.  Benedict's  comparison  of  their  shape  to  that  of  a  retort  is  excel- 
lent. The  ovary  is  early  decadent.  The  relations  of  the  organs  of  the 
interovarial  space  are  typical  of  the  genus  as  Benedict  has  described 
them.  The  vitellaria  are  follicular.  The  follicles  are  not  large  nor  are 
they  closely  packed.  They  too  are  decadent  early.  The  uterus  (Figs. 
51,  52)  in  ripe  proglottids  consists  of  a  median  tube  and  9-14  lateral 
pouches  on  either  side.  These  pouches  occupy  the  ventral  part  of  the 
segment  while  the  testes  are  dorsal.  *'The  eggs  which  are  found  in 
the  uterus  have  a  diameter  of  0.02  mm."  (Benedict).  Measurements 
by  the  writer  gave  a  range  of  0.019-0.021  mm.  for  the  embryo  and 
0.036-0.046  mm.  for  the  second  membrane.  The  outer  membrane  meas- 
ures 0.038-0.060  mm.  The  second  membrane  is  thick  and  granular  while 
the  outer  membrane  in  uterine  eggs  is  thick  and  hyaline. 

This  species  somewhat  resembles  P.  piisillus  Ward  but  is  differenti- 
ated from  that  species  by  the  smaller  size  of  its  head,  by  its  very  much 
smaller  suckers  and  smaller  sucker  openings.  This  species  has  a  much 
longer  neck  than  has  P.  pusillus.  The  character  of  the  segmentation  of 
the  two  worms  is  utterly  different  being  very  evident  in  P.  pusillus^ 
with  deep  constrictions  between  proglottids.  In  P.  exiguus  segmenta- 
tion is  not  evident.    The  number  of  proglottids  in  P.  pusillus  is  greater 


58  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [58 

than  in  P.  exiguus.  The  mass  of  coils  of  the  vas  deferens  in  P.  pusillus 
is  much  smaller  than  in  P.  exiguus  and  it  lies  almost  wholly  anterior 
to  the  cirrus-pouch  while  in  P.  exiguus  it  lies  more  nearly  at  the  end  of 
the  cirrus-pouch.  The  testes  are  more  numerous  in  P.  pusillus  and  the 
cirrus-pouch  is  much  shorter  than  in  P.  exiguus.  The  lobes  of  the 
ovaries  in  P.  pusillus  are  bent  strongly  posteriad  while  such  a  condition 
is  never  so  marked  in  P.  exiguus.  The  proportions  of  the  proglottids 
are  greatly  different. 

P.  exiguus  closely  resembles  P.  fallax  La  Rue.  It  differs  from  that 
species  in  having  a  smaller  head,  smaller  suckers,  a  smaller  fifth  sucker, 
a  shorter  neck,  fewer  and  smaller  proglottids,  more  testes,  a  shorter 
cirrus  when  evaginated,  and  a  greater  number  of  lateral  uterine  out- 
pocketings.  The  embryos  of  P.  exiguus  are  smaller  than  in  P.  fallax. 
The  differences  are  of  such  a  character  and  of  such  a  constancy  as  to 
be  of  specific  importance.  This  species  resembles  P.  a^gonis  Barbieri 
even  more  than  it  does  P.  fallax  except  that  P.  agonis  has  no  well  de- 
veloped fifth  sucker.  In  total  length  and  in  the  size  of  the  proglottids 
P.  exiguus  is  the  larger.  P.  exiguus  has  more  uterine  pouches  and 
fewer  but  larger  testes  than  has  P.  agonis.  Unfortunately  Barbieri 's 
description  fails  in  some  particulars,  nevertheless  sufficient  data  are 
given  to  establish  the  difference  between  the  two  species.  P.  exiguus 
resembles  P.  duhius  La  Rue  but  its  head,  its  suckers,  its  proglottids, 
and  its  cirrus-pouch  are  smaller.  These  two  species  have  about  the 
same  number  of  testes.  The  embryo  of  P.  dubius  is  much  larger  than 
that  of  P.  exiguus.  P.  exiguus  differs  radically  from  P.  ambiguus  and 
P.  esocis  in  the  possession  of  a  fifth  sucker.  Other  points  of  difference 
are  also  to  be  found.  P.  exiguus  is  smaller  than  other  North  American 
species  of  Proteocephaliis. 

PROTEOCEPHALUS  PUSILLUS  Ward 

[Figs.  2-4,  53-55] 

1910 :  Proteocephalus  pusillus  Ward,  1910 :1185-1187 
1911:  Proteocephalus  pusillus  La  Rue,  1911:475 
Specific  Diagnosis:  Characters  of  the  genus.  Cestodes  small, 
length  30-50  mm.  Maximum  breadth  0.350  mm.  Proglottids  few.  Seg- 
mentation distinct.  Head  spheroidal,  frequently  much  contracted, 
breadth  about  0.300  mm.,  length  about  0.260  mm.  Four  suckers  placed 
at  broadest  zone  of  head.  Suckers  deep,  0.140  mm.  long  by  0.110  mm. 
broad.  Fifth  sucker  well  developed,  muscular,  0.060  mm.  in  diameter. 
Neck  1.0-1.5  mm.  long  by  0.21  mm.  broad.  First  proglottids  broader 
than  long,  breadth  of  same  0.090  mm.    Mature  proglottids  longer  than 


59]  PROTEOCEPHALIDAE  —  LA  RUE  » 

broad.  Eipe  proglottids  much  longer  than  broad.  Length  of  same 
0.84-1.4  mm.,  breadth  0.18-0.35  mm.    End-proglottid  present  and  fertile. 

Genital  sinus  marginal,  irregularly  alternating,  situated  at  end  of 
first  %-%  of  proglottid.  Vagina  usually  anterior  and  dorsal  to  cirrus- 
pouch,  rarely  posterior.  Testes  44-60-70  in  number,  arranged  in  two 
layers  between  vitellaria.  Length  of  same  0.06-0.096  mm.,  breadth 
0.04-0.05  mm.  Vas  deferens  a  small  mass  of  coils  anterior  to  cirrus- 
pouch.  Cirrus-pouch  0.095-0.106  mm.  long  by  0.053-0.060  mm.  broad, 
reaching  5^-14  across  the  proglottid  breadth.  Ductus  ejaculatorius 
forming  one  or  two  coils  in  cirrus-pouch.  Cirrus  short  and  straight. 
Vagina  never  crossing  cirrus-pouch.  Sphincter  vaginae  and  receptacu- 
lum  seminis  not  seen.  Ovary  bilobed,  posterior.  Lobes  short,  thick, 
free  ends  of  same  frequently  pressed  posteriad  and  together.  Vitellaria 
sparse,  follicles  small.  Uterus  with  10-14-16  lateral  pouches  on  either 
side.    Eggs  not  observed. 

Habitat:  Intestine  and  esophagus  of  Salmo  sehago  Girard  (type 
host),  Sebago  Lake,  Me.  (type  locality)  ;  intestine  and  pyloric  coeca  of 
Cristivomer  namaycush  Walbaum  ( ?),  Lake  Temagami,  Ontario. 

Type:  Material  in  bottles  15,  16,  42,  in  Professor  "Ward's  collec- 
tion from  Sebago  Lake,  Me.,  summer  of  1907,  and  slides  from  this 
material. 

"Ward  (1910:1185-1187)  first  described  this  species  which  he  had 
found  in  the  Sebago  salmon.    "Ward's  description  follows. 

"Proteocephalus  pusUlus  nov.  spec. — Adult  cestode  with  short  strobila,  meas- 
uring only  30  to  50  mm.  in  length,  Proglottids  scanty,  segmentation  distinct. 
Head  much  contracted.  Neck  i  to  1.5  mm.  long  by  0.21  mm.  broad.  First  pro- 
glottids 0.09  mm.  broad  changing  gradually  until  in  mature  proglottids  the  length 
greatly  exceeds  the  breadth.  Ripe  proglottids  measure  0.84  to  1.4  mm.  long  by  0.18 
to  0.35  mm.  broad.  Terminal  proglottids  present  and  fertile.  Sexual  organs 
typical  for  Proteocephalus ;  uterus  median,  with  10  to  14  lateral  outpocketings  on 
either  side.  Testes  numerous,  within  vitellaria.  Genital  pore  lateral,  one-third  to 
two-fifths  of  length  of  proglottid  from  anterior  margin  of  same.  Ovaries  bilobed, 
median  isthmus  indistinct,  anteroposterior  diameter  nearly  equal  to  breadth  of 
both  lobes.  Only  a  few  specimens  obtained  from  a  single  host  species,  Salmo 
sehago. 

"This  species  approaches  most  nearly  to  P.  ocellata  and  P.  percae  among 
known  species.  Unlike  the  new  species,  however,  both  of  these  older  forms  have 
a  fifth  sucker,  a  few  lateral  uterine  outpocketings,  a  longer  neck,  differently  shaped 
ovaries,  and  markedly  different  proglottids. 

"In  specimens  with  developed  proglottids  the  head  was  so  much  contracted 
or  distorted  that  any  special  description  would  be  of  little  value.  One  could 
easily  observe  the  general  features  characteristic  of  the  genus.  There  was  no 
well  developed  terminal  or  fifth  sucker,  and  the  end  organ,  which  is  known  to 


€0  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [60 

replace  it  in  many  forms  of  this  genus,  was  inconspicuously  developed,  if  present. 
Personally,  1  incline  to  the  view  that  on  more  careful  examination  this  structure 
will  be  found  in  all  species,  even  those  in  which  its  absence  has  been  made  a 
matter  of  record.  Accordingly,  not  much  weight  can  be  put  in  its  presence  or 
absence  in  any  individual. 

"Three  plerocercoid  larvae  or  young  cestodes  were  found  in  company  with 
Proteocephalus  pusillus,  which  I  regard  as  young  forms  of  this  species.  The 
largest  came  from  the  salmon  which  was  most  heavily  infected  with  this  cestode 
parasite.  It  was  3.15  mm.  long  and  had  begun  to  assume  clearly  the  appearance 
of  an  immature  cestode.  The  head  measured  0.3  mm.  wide  by  0.26  mm.  long,  and 
the  suckers  0.14  mm.  in  length  by  o.ii  mm.  in  width.  The  neck  was  slightly  nar- 
rower than  the  head,  but  was  not  clearly  set  off  from  the  body,  which  was  very 
imiform  in  diameter  and  measured  0.25  mm.  in  average  width.  The  posterior  end 
of  the  body  was  swollen  into  a  rounded  knob  about  0.35  mm.  broad  and  of  ap- 
proximately the  same  length.  This  feature  was  evidently  produced  by  a  powerful 
contraction  of  the  terminal  region  of  the  body.  In  and  near  it  one  could  see  very 
indistinct  indications  of  proglottid  formation.  In  form,  size,  and  general  aspect 
this  young  cestode  was  in  full  agreement  with  the  anterior  regions  of  the  mature 
cestodes  of  this  species  with  which  it  was  associated.  The  head,  which  was  not 
contracted,  showed  on  careful  study  the  delicate  outline  of  a  rudimentary  end 
organ.  While  such  a  structure  was  not  demonstrated  in  the  mature  individuals 
described  above,  one  can  say  positively  that  if  present  it  could  not  have  been  seen 
owing  to  the  greatly  contracted  condition  of  the  adult  scolices.  I  believe  that  its 
presence  will  be  demonstrated  in  more  favorable  specimens.  The  complete  agree- 
ment of  this  largest  larva  with  the  mature  specimens  in  all  other  features  compels 
me  to  regard  both  as  different  stages  in  the  development  of  the  same  species. 

"The  other  larvae  were  still  in  early  stages  of  development  and  probably  had 
been  ingested  by  the  salmon  at  a  very  recent  date.  Their  relationship  is  not  so 
clear  in  all  respects,  and  yet  I  do  not  hesitate  to  associate  with  the  new  species  of 
Proteocephalus  a  plerocercoid  or  young  cestode  obtained  from  the  same  host  as 
the  adult  worms  and  the  older  larva  just  described.  The  head  is  broadly  conical, 
without  furrows,  and  measures  0.3  mm.  in  breadth.  The  suckers  measure  60  to 
74  fjL.  in  diameter.  There  is  no  rostellum  or  fifth  sucker  to  be  found,  while  the  end 
organ  is  so  poorly  developed  as  to  be  visible  with  difficulty  and  only  under  the 
most  favorable  circumstances.  The  neck  is  nearly  as  broad  as  the  head.  In  gen- 
eral appearance  this  larva  resembles  the  adult  cestode  and  the  older  larva  pre- 
viously described.  With  some  reserve  one  may  also  assign  to  this  species  a  single 
plerocercus  taken  from  another  specimen  of  Sabno  sebago.  The  head,  which  meas- 
ures only  150  ^  in  breadth,  is  shaped  like  that  of  the  young  cestode  and  like  it  is 
without  rostellum  or  fifth  sucker,  while  the  end  organ  is  difficult  to  demonstrate. 
Neither  furrows  nor  ridges  are  seen  on  the  larva,  which  has  a  total  length  of  1.14 
mm.  The  sucker  measures  only  30  to  45  fi  in  diameter.  The  neck  is  slightly  nar- 
rower than  the  head.  This  form  certainly  belongs  to  the  genus  Proteocephalus 
and  probably  to  the  species  already  described." 


61]  PROTEOCEPHALIDAE  —  LA  RUE  61 

La  Rue  (1911:475)  listed  this  form  among  other  Proteocephalus 
species.  Dr.  H.  B.  Ward  has  very  kindly  loaned  the  writer  his  speci- 
mens for  study.  More  preparations  have  been  made  and  from  them 
the  writer  has  been  able  to  secure  additional  data  on  this  species. 

The  type  material  in  bottles  15,  16  and  42  was  taken  from  the 
intestine  and  oesophagus  of  Salmo  sehago  Girard,  Sebago  Lake,  Me.  Jy. 
27  and  Aug.  6,  1907.  Other  specimens  which  the  writer  has  assigned 
to  this  same  species  were  taken  by  Dr.  H.  B.  Ward  from  two  specimens 
of  Cristivomer  namaycush  Walbaum  (?),  Lake  Temagami,  Ontario,  in 
August,  1911.    These  last  are  described  separately. 

This  is  one  of  the  smaller  species  of  Proteocephalus.  The  observed 
length  is  from  30-50  mm.  for  specimens  which  have  ripe  proglottids. 
Proglottids  are  few  in  number  and  the  segmentation  is  distinct.  The 
head  is  spheroidal  but  in  many  cases  it  is  so  greatly  contracted  that  its 
structure  can  not  be  determined.  The  head  measures  0.300  mm.  broad 
by  about  0.260  mm.  long.  The  four  suckers  measure  about  0.140  mm. 
long  by  0.110  mm.  broad.  The  sucker  cavity  is  quite  deep.  At  the 
apex  of  the  head  (Figs.  2,  3,  4)  is  a  fairly  well  developed  fifth  sucker 
which  in  well  stained  specimens  shows  an  evident  musculature  and  a 
cavity.  Ward  (1910)  reported  only  an  end  organ  or  vestigial  fifth 
sucker  instead  of  a  fifth  sucker.  When  his  specimen  was  restained  and 
mounted  it  showed  a  well  formed  fifth  sucker.  Characteristic  fifth 
suckers  measure  about  0.060  mm.  in  diameter.  The  neck  is  1-1.5  mm. 
long  by  about  0.21  mm.  broad.  It  passes  over  quickly  into  the  first  pro- 
glottids which  are  broader  than  long.  Their  breadth  is  about  0.090  mm. 
The  proglottids  rapidly  become  quadrate  or  even  longer  than  broad. 
Ripe  proglottids  measure  0.84-1.4  mm.  long  by  0.18-0.35  mm.  broad.  An 
end-proglottid  is  usually  present  and  fertile. 

The  common  genital  sinus  is  marginal,  situated  at  the  end  of  the 
1/  first  %-%  of  the  proglottid.  Its  position  alternates  irregularly.  The 
testes  (Figs.  54,  55)  number  44-60-70.  Since  these  lie  in  two 
layers  the  figure  does  not  show  all  of  them.  In  less  mature  proglottids 
where  the  testes  are  much  smaller  and  more  distinct  they  can  be  counted 
more  readily.  The  testes  measure  0.06-0.072-0.096  mm.  long  by  0.040- 
0.050  mm.  broad.  The  vas  deferens  (Figs,  53,  54,  55)  forms  a  small 
mass  of  coils  in  the  middle  of  the  proglottid.  The  larger  part  of  the 
mass  is  anterior  to  the  cirrus-pouch,  a  condition  not  usually  met  with 
in  this  genus.  The  cirrus-pouch  which  is  rather  heavy  reaches  34 -H. 
across  the  proglottid  breadth.  Its  length  is  0.095-0.106  mm.  and  its 
breadth  0.053-0,060  mm.  The  ductus  ejaculatorius  forms  a  coil  or  two 
within  the  cirrus-pouch  before  passing  over  into  the  short  straight  cir- 
rus which  has  not  been  seen  protruded.    On  account  of  a  lack  of  suffi- 


62  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [62 

cient  material  but  very  few  sections  have  been  made  and  these  were  too 
poor  to  yield  much  valuable  data. 

The  vagina  (Figs.  54,  55)  usually  opens  anterior  and  dorsal  to  the 
cirrus-pouch  but  rarely  (Fig.  53)  it  is  posterior  to  the  same.  The 
number  of  proglottids  examined  have  been  too  few  to  establish  whether 
the  posterior  position  of  the  vagina  is  normal.  Even  when  in  the  ante- 
rior situation  the  vagina  does  not  cross  the  cirrus-pouch.  From  its 
opening  it  makes  a  long  curve  toward  the  middle  of  the  proglottid 
which  it  reaches  at  a  point  posterior  to  the  inner  end  of  the  cirrus- 
pouch.  The  vagina  is  dorsal  to  the  uterus.  A  vaginal  sphincter  could 
not  be  demonstrated,  and  if  present  it  must  be  very  weak.  The  pres- 
ence of  a  receptaculum  seminis  has  not  been  demonstrated.  The  bilobed 
ovary  (Figs.  53,  54,  55)  is  peculiar  in  that  the  lobes  which  are  short  and 
very  thick  are  pressed  closely  together.  This  is  a  condition  not  yet 
found  regularly  in  any  other  species  of  the  genus  altho  Nufer  (1905) 
found  it  in  P.  macrocephalus.  In  transverse  sections  the  thickness  and 
compactness  of  the  ovary  are  noted  at  once.  The  vitellaria  are  sparse, 
the  follicles  small.  The  ducts  of  the  interovarial  space  have  not  been 
thoroly  investigated.    An  ootype  and  an  oocapt  are  present. 

In  ripe  ploglottids  the  uterus  (Fig.  53)  shows  10-14-16  lateral  out- 
pocketings  on  either  side.  No  uterine  pores  have  been  seen.  The  eggs 
have  not  been  observed. 

Other  specimens  which  the  writer  has  provisionally  assigned  to  this 
species  were  taken  by  Dr.  H.  B.  Ward  from  the  pyloric  and  intestinal 
regions  of  Cristivomer  namaycush  Walbaum  (?),  Lake  Temagami,  On- 
tario, August,  1910.  It  seems  that  the  determination  of  the  host  must 
remain  somewhat  in  doubt -f or  there  was  no  attempt  at  the  identification 
of  the  fish  when  it  was  caught  and  it  was  not  preserved.  The  identifica- 
tion was  made  from  the  memory  of  the  appearance  of  the  fish  after  the 
return  of  the  expedition.  Both  Professor  Reighard  and  Professor 
Ward  when  questioned  by  the  writer  thought  there  could  be  little  doubt 
of  the  determination.  These  specimens  bear  the  lot  numbers  Tip,  Tlj, 
T2h,  and  T2p  in  Dr.  H.  B.  Ward's  collection. 

The  specimens  are  small,  measuring  up  to  20-30  mm.  long.  No 
specimens  have  ripe  proglottids.  When  the  worm  is  fully  mature  it  is 
perhaps  considerably  longer.  Measurements  of  a  number  of  specimens 
yielded  the  following  data :  Average  breadth  of  seven  heads  0.284  mm., 
maximum  breadth  of  same  0.340  mm.,  minimum  0.244  mm.;  average 
length  of  six  heads  0.177  mm.,  maximum  length  of  same  0.238  mm.,  min- 
imum 0.133  mm. ;  average  length  of  seventeen  suckers  0.134  mm.,  maxi- 
mum length  of  same  0.179  mm.,  minimum  0.106  mm.;  average  breadth 
of  thirteen  suckers  0.108  mm.,  maximum  breadth  of  same  0.127  mm., 


63]  PROTEOCEPHALIDAE  —  LA  RUE  63 

minimum  0.096  mm. ;  sucker  opening  0.058-0.080  mm. ;  diameter  of  fifth 
sucker  about  0.060  mm.  Average  length  of  six  necks  1.49  mm.,  maxi- 
mum length  1.70  mm.,  minimum  1.02  mm.;  neck  narrow,  up  to  0.135 
mm.  The  proglottids  are  few,  numbering  about  30-40.  First  proglot- 
tids  are  broader  than  long,  or  quadrate,  length  soon  exceeds  breadth, 
and  mature  proglottids  are  longer  than  broad  measuring  up  to  0.56-0.68 
mm.  long  by  0.24-0.265  mm.  broad.  The  maximum  breadth  of  0.340 
mm.  was  observed  in  a  proglottid  measuring  0.357  mm.  long  by  0.340 
mm.  broad.    Segmentation  is  very  distinct. 

The  genital  pore  is  near  the  middle,  usually  anterior  thereto  but 
sometimes  posterior.  The  vagina  lies  anterior  to  the  cirrus-pouch. 
There  are  about  44-66  testes  and  these  measure  about  0.159  by  0.180 
mm.  in  diameter.  They  form  two  irregular  layers  in  the  region  bounded 
by  the  vitellaria  and  ovary.  In  many  proglottids  of  this  lot  the  testes 
are  not  as  fully  developed  as  in  the  proglottids  of  specimens  from  Salmo 
sehago.  This  condition  facilitates  the  counting  of  the  testes.  The  coils 
of  the  vas  deferens  form  a  small  mass  situated  for  the  most  part  anterior 
to  the  cirrus-pouch.  Within  the  cirrus-pouch  the  ductus  ejaculatorius 
forms  a  single  coil  and  then  it  passes  over  into  the  small  cirrus.  The 
cirrus-pouch  usually  extends  to  the  middle  of  the  proglottid  or  nearly 
thereto.  It  has  a  length  of  about  0.111-0.150  mm.  The  average  length 
of  eight  pouches  was  0.133  mm.  In  breadth  the  cirrus-pouch  measures 
0.053-0.070  mm.  The  lobes  of  the  ovary  are  thick  and  are  bent  back 
until  they  nearly  meet  in  the  median  line.  Vitellaria  are  sparse.  A 
fully  developed  uterus  has  not  been  observed.  Of  the  drawings  repre- 
senting P.  pusillus,  figures  2,  4,  and  55  were  made  from  specimens  which 
came  from  Cristivomer  namaycush.  All  the  other  drawings  of  this  spe- 
cies were  made  from  specimens  taken  from  Salmo  sehago. 

In  general  these  two  forms  agree  very  well  but  there  are  two  points 
of  minor  disagreement.  These  are  in  regard  to  the  length  of  the  cirrus- 
pouch  and  the  size  of  the  suckers.  As  to  the  former  the  difference  in 
length  is  but  a  matter  of  a  few  micra  and  this  perhaps  may  be  explained 
by  the  fact  that  the  proglottids  are  in  different  states  of  contraction. 
As  for  the  other  point  of  difference  it  may  be  stated  that  in  the  speci- 
mens taken  from  Cristivomer  namaycush  the  suckers  are  on  the  average 
about  the  size  of  the  suckers  of  the  typical  P.  pusillus  but  in  one  or  two 
cases  observed  the  length  and  breadth  of  the  sucker  is  appreciably 
larger.  It  is  to  be  remembered  that  the  measurements  of  the  suckers 
of  the  typical  P.  pusillus  are  based  on  a  single  head.  Consequently  the 
range  of  size  of  suckers  in  that  species  has  not  been  determined.  Tho 
no  ripe  proglottids  were  present  in  the  material  from  Cristivomer  no- 


64  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [64 

maycush  and  consequently  some  of  the  best  diagnostic  features  could 
not  be  determined  it  seems  best  to  consider  these  forms  identical. 

Proteocephalus  pusUlus  Ward  is  closely  allied  to  P.  exiguus  La  Rue 
but  is  differentiated  from  that  species  by  its  larger  head,  larger  suckers, 
and  larger  sucker  openings.  The  measurements  of  proglottids  of  the 
two  species  are  much  alike.  However,  the  type  of  segmentation  is  very 
dissimilar.  In  the  reproductive  organs  P.  pusillus  varies  from  P.  exi- 
guus in  having  more  numerous  and  larger  testes.  The  testes  of  P^ 
pjisillus  lie  in  two  layers  while  in  P.  exiguus  they  are  in  one  layer.  The 
cirrus-pouch  of  P.  exiguus  is  much  longer  than  that  of  P.  pusillus.  The 
ovaries  of  the  two  species  are  very  different.  In  the  one  species  the 
lobes  are  slightly  arched  while  in  the  other  the  lobes  are  bent  nearly 
together.  The  lateral  uterine  pouches  of  P.  exiguus  are  larger  and 
scarcely  as  numerous  as  in  P.  pusillus.  P.  pusillus  somewhat  resembles 
P.  fallax  La  Rue  but  it  is  smaller.  It  has  a  larger  head  and  suckers, 
more  numerous  and  larger  testes,  a  much  shorter  cirrus-pouch,  and 
more  numerous  uterine  outpocketings  than  P.  fallax.  The  ovaries  of 
the  two  species  are  very  different. 

P.  pusillus  differs  from  P.  agonis  Barbieri  in  having  a  fifth  sucker. 
Moreover  P.  pusillus  is  a  larger  form  than  P.  agonis.  P.  agonis  has 
smaller  and  more  numerous  testes  than  P.  pusillus,  and  also  fewer 
uterine  outpocketings.  P.  pusillus  differs  radically  in  size  and  propor- 
tions from  P.  percae  (0.  F.  Miiller).  P.  pusillus  differs  from  P.  esocis 
(Schneider)  in  having  a  larger  head,  larger  suckers,  in  possessing  a 
fifth  sucker,  and  a  shorter  neck.  The  proportions  of  the  segments  are 
different.  The  length  of  the  cirrus-pouch  and  the  ratio  of  its  length  to 
the  proglottid  breadth  differ  greatly  in  the  two  species.  The  character 
of  the  segmentation  is  very  unlike.  There  is  very  little  probability  that 
P.  pusillus  is  the  same  as  P.  salvelini  (Linton).  P.  salvelini  has  no  fifth 
sucker.  Its  head,  suckers,  and  cirrus-pouch  are  larger  than  those  of 
P.  pusillus.  Linton's  description  of  P.  salvelini  does  not  furnish  many 
characters  for  determination  and  his  drawing  of  the  proglottid  is  so 
indistinct  that  it  cannot  be  relied  upon  as  a  source  of  data. 


65] 


PROTEOCEPHAUDAE  —  LA  RUE 


65 


PROTEOCEPHALUS  PINGUIS  La  Rue 
[Figs.  29-32,  35,  36,  88-93] 
1911 :    Proteocephalus  pinguis  La  Rue,  1911 :478. 

Specific  Diagnosis:  Characters  of  genus.  Strobila  short  and  slen- 
der. Observed  length  up  to  90  mm.  Maximum  breadth  1.24  mm.  Neck 
3-7  ram.  long  by  0.20-0.25  mm.  broad.  Proglottids,  first  in  chain  very 
short,  0.05  mm.  long  by  0.25  mm.  broad.  Mature  and  ripe  proglottids 
nearly  quadrate  or  in  a  few  ripe  proglottids  length  exceeding  the 
breadth,  0.595  by  0.595  mm.  up  to  0.730  mm.  long  by  0.595  mm.  broad. 
End-proglottid  present  and  functional.  Segmentation  not  very  distinct. 
Intersegmental  furrows  shallow.  Head  conical,  flattened  dorso-ventrally, 
without  furrows,  presenting  great  variations  in  shape.  Breadth  of  head 
about  0.33  mm.,  thickness  0.22  mm.,  length  0.20-0.25  mm.  Suckers  cup- 
shaped,  deep,  muscular.  Diameter  of  suckers  0.095-0.105  mm.,  of  aper- 
ture of  sucker  0.06-0.07  mm.    Diameter  of  fifth  sucker  0.05-0.075  mm. 

Common  genital  sinus  irregularly  alternating,  situate  at  or  near 
middle  of  lateral  margin  of  proglottid.  Testes  ovoidal,  0.05  mm.  long 
by  0.04-0.05  mm.  broad,  54-70  in  number,  in  a  single  layer  taking  up 
field  between  vitellaria  anterior  to  ovary.  Ductus  ejaculatorius  nearly 
straight.  Cirrus  well  muscled,  straight,  length  when  protruded  0.07- 
0.08  mm.  by  0.02-0.025  mm.  broad.  Cirrus-pouch  short,  stout,  0.13-0.14 
mm.  long  by  0.05-0.06  mm.  broad.  Ratio  of  length  of  cirrus-pouch  to 
breadth  of  proglottid  1 :3  or  1 :4.  Vagina  anterior,  but  vaginal  opening 
always  dorsal  to  cirrus-pouch.  Vagina  crossing  inner  end  of  cirrus- 
sheath.  Sphincter  vaginae  small.  Receptaculum  seminis  small,  situate 
anterior  to  ovary.  Uterus,  when  fully  developed,  with  10-14  lateral 
pouches  on  either  side.  Ventral  uterine  pores  2-3  in  number.  Embryos 
0.016-0.018  mm.  in  diameter. 

Habitat:    In  intestine  of  host. 


Host 

Locality 
Sebago  Lake, 

Collector 

Authority 

Esox  reticulatus 

H.  B.  Ward 

La  Rue 

Le  Sueur 

Maine 

(the  present  paper ; 

Esox  lucius 

"Walnut  Lake, 

T.  L.  Hankinson 

La  Rue 

Linn. 

Mich. 

(the  present  paper) 

Esox  lucius 

Lake  Geneva, 

A.  J.  Coats 

La  Rue 

Linn. 

Wis. 

(the  present  paper) 

Type:     Alcoholics  in  Dr.  H.  B.  Ward's  Sebago  Lake  collection 
Nos.  4,  6,  7,  8,  9,  107.    Slides  of  same.    The  material  was  collected  by 


66  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [66 

Dr.  H.  B.  Ward  from  Esox  reticulatus  Le  Sueur  (type  host) ;  Sebago 
Lake,  Maine  (type  locality). 

Co-type:  Alcoholics  No.  TLH413  in  Dr.  H.  B.  Ward's  collection, 
from  Esox  lucius  Linn.,  Walnut  Lake,  Michigan,  and  slides  of  the  same. 

Autotyi)e :  Alcoholics  No.  10.35  in  collection  Dr.  H.  B.  Ward  Irom 
Esox  lucius  Linn.,  Lake  Greneva,  Wisconsin. 

Leidy  (1888:169  and  1890:417)  described  some  specimens  of  ces- 
todes  from  Esox  reticulatus.  The  two  specific  names  which  he  proposed 
for  them  the  writer  considers  to  be  synonyms  and  has  included  under 
the  one  name,  Proteocephalus  nematosomu  (Leidy).  Leidy 's  specimens 
were  considerably  larger  than  the  specimens  of  P.  pinguis  and  appar- 
ently are  not  the  same.  A  more  complete  discussion  of  Leidy 's  species 
occurs  later  in  the  article.  La  Rue  (1911:478)  briefly  characterized 
this  species  and  proposed  for  it  the  name  Proteocephalus  pinguis. 

This  study  is  based,  for  the  most  part,  upon  specimens  collected  by 
Dr.  H.  B.  Ward  from  the  intestines  of  Esox  reticulatus  Le  Sueur  at 
Sebago  Lake,  Maine,  while  engaged  in  an  investigation  of  that  lake  for 
the  U.  S.  Bureau  of  Fisheries,  in  the  summer  of  1907.  Portions  of  the 
study  are  also  based  upon  specimens  secured  by  Prof.  T.  L.  Hankinson 
from  Esox  lucius  Linn,  in  the  course  of  a  biological  investigation  of 
Walnut  Lake,  Mich.,  during  the  summer  of  1906.  This  last  material 
was  referred  to  in  a  former  paper.  La  Rue  (1909:25),  as  "A  new  spe- 
cies, which  I  shall  describe  in  detail  later ".    It  bears  the  number 

TLH413  in  Professor  Ward's  collection.  Dr.  H.  B.  Ward  examined  six 
specimens  of  Esox  reticulatus  Le  Sueur  while  at  Sebago  Lake.  These 
six  fish  yielded  sixteen  entire  Proteocephalids  and  pieces  as  follows: 
No.  4,  1  head-l-2  pieces;  No.  6,  2  heads+2-3  pieces;  No.  7,  1  head-|-2 
pieces;  No.  8,  11  headsH-2  pieces;  No.  9,  1  head;  No.  107,  5  heads,  1 
young  cestode-f-10  large  pieces.  In  many  instances  the  specimens  re- 
ported as  heads  were  complete  worms  while  in  others  the  end  proglottid 
and  part  of  the  strobila  were  missing.  Thus  the  infection  of  Sebago 
Lake  hosts  was  light.  Hankinson 's  material  in  bottle  No.  413  from 
Esox  lucius  of  Walnut  Lake  was  probably  from  a  single  host.  It  con- 
tained 60-70  si)ecimens  of  P.  pinguis.  A  bottle,  No.  10.35  in  Dr.  H.  B. 
Ward's  collection,  bearing  the  label  "from  stomach  and  intestine  of  a 
'pickerel,'  Lake  Geneva,  Wisconsin,  Jy.  1,  1904.  A.  J.  Coats,  collector", 
contained  5  or  6  complete  specimens  of  P.  pinguis  La  Rue  and  several 
pieces  of  the  same  species.  Replying  to  a  letter  of  inquiry  Prof.  (Jeoi^ 
Wagner  of  the  University  of  Wisconsin  states  that  only  one  species  of 
pickerel,  the  Esox  lucius,  is  found  in  Lake  (Jeneva,  Wisconsin.  The  data 
on  this  species  were  obtained  from  specimens  cleared  and  examined  in 


67]  PROTEOCEPHALIDAE  —  LA  RUE  67 

glycerine,  from  specimens  stained  and  mounted  in  toto  and  from  trans- 
verse and  frontal  sections. 

These  cestodes  are  small  and  slender  (Fig.  32).  The  largest  speci- 
men from  the  Sebago  Lake  material  measured  90  mm.  long  by  1.24  mm. 
in  maximum  breadth.  The  longest  specimen  from  the  Walnut  Lake  host 
measured  66  mm.  long  by  1.0  mm.  in  maximum  breadth.  The  strobila  of  a 
50  mm.  specimen  is  made  up  of  an  unsegmented  neck  3-7  mm.  long  by 
0.20-0.25  mm.  broad  followed  by  300  proglottids.  The  youngest  proglot- 
tids  measure  about  0.050  mm.  long  by  0.250  mm.  broad.  Further  along 
in  the  chain  these  increase  in  length  and  breadth  until  in  mature  and 
ripe  proglottids  the  length  and  breadth  are  about  equal.  Measurements 
of  several  ripe  proglottids  are  as  follows,  the  length  in  millimeters  being 
stated  first:  0.595  by  0.595,  0.680  by  0.680,  0.595  by  0.663,  0.730  by 
0.595  mm.    In  transection  the  proglottids  are  elliptical. 

A  typical  functional  end-proglottid  with  a  rounded  posterior  end 
is  present.  The  segmentation  is  not  very  plain,  for  the  inter-segmental 
furrows  are  not  deep  and  the  corners  of  the  proglottids  do  not  project. 
The  lateral  margins  are  rounded  and  smooth.  The  head  (Figs.  29-32, 
35,  36)  is  a  cone-shaped  structure  somewhat  flattened  dorso-ventrally.  It 
is  not  marked  by  folds  or  furrows.  Its  breadth  is  about  0.33  mm.,  its 
thickness  about  0.22  mm.,  its  length  about  0.20-0.25  mm.  In  extreme 
cases  the  head  may  attain  a  breadth  of  0.45  mm.  This  extreme  breadth 
is  due  to  a  contraction  of  the  longitudinal  muscles  of  the  head  and  neck. 
A  collection  of  these  specimens  shows  heads  of  very  different  states  of 
contraction.  The  head  bears  on  its  broadest  part  four  deep  cup-shaped 
muscular  suckers  which  are  directed  forward  and  outward.  On  its  apex 
the  head  bears  a  well  formed  muscular  fifth  sucker.  The  diameter  of 
the  suckers  ranges  from  0.095-0.105  mm.,  while  the  diameter  of  the 
aperture  varies  from  0.06-0.07  mm.  The  fifth  sucker  measures  0.05-0.075 
mm.  in  diameter.  Altho  its  cavity  is  quite  shallow  it  has  the  appearance 
of  being  functional  for  it  possesses  a  typical  musculature  (Fig.  93). 

The  cuticula  and  musculature  have  not  been  found  to  differ  in  their 
essential  features  from  those  of  other  members  of  the  genus.  The  ner- 
vous system  in  the  head  is  much  like  that  of  Ophiotaenia  filaroides.  A 
ganglionic  mass  is  located  at  the  level  of  the  suckers  (Fig.  93).  Two 
main  lateral  nerve  trunks  extend  throughout  the  length  of  the  strobila. 
No  accessory  nerve  trunks  were  observed. 

The  excretory  system  in  the  head  is  made  up  of  four  main  longi- 
tudinal canals  which  at  various  levels  receive  many  anastomosing  coiled 
vessels  which  pass  through  the  tissues  of  the  scolex  and  neck  in  every 
direction.  In  the  strobila  there  are  two  pairs  of  lateral  excretory  ves- 
sels, ventral  and  dorsal  respectively.     These  lie  within  the  medullary 


68  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [68 

parenchyma.  The  dorsal  vessel  passes  dorsad  and  the  ventral  vessel 
ventrad  to  the  cirrus-pouch  and  vagina.  In  diameter  the  ventral  vessel 
greatly  exceeds  the  dorsal  and  its  lining  membrane  is  thin  while  the 
membrane  lining  the  dorsal  vessel  is  thick.  A  posterior  excretory  com- 
missure in  each  proglottid  could  not  be  found.  Numerous  secondary 
excretory  openings  which  are  connected  with  the  ventral  vessel  and  less 
frequently  with  the  dorsal  vessel  have  been  seen.     The  drawing  (Fig. 

91)  shows  rather  an  extraordinary  case  in  that  the  duct  is  larger  and 
more  extensive  in  its  connections  than  is  usual.  In  the  end-proglottid 
the  four  main  excretory  ducts  discharge  to  the  exterior  by  a  common 
pore.    A  bladder  could  not  be  seen. 

The  common  genital  sinus  is  marginal,  situated  at  or  near  the  mid- 
dle of  the  proglottid.  While  the  vagina  lies  anterior  to  the  cirrus-pouch, 
the  opening  of  the  vagina,  without  exception,  is  dorsal  to  the  opening  of 
the  cirrus  (Figs.  88,  90).    There  is  no  genital  papilla.    The  testes  (Fig. 

92)  are  numerous,  54-70.  These  are  in  a  single  layer  in  the  medullary 
parenchyma,  occupying  nearly  the  entire  field  between  the  vitellaria 
anterior  to  the  ovary.  The  testes  measure  as  much  as  0.05  mm.  long  by 
0.04-0.05  mm.  broad.  The  vasa  efferentia  (Fig.  90)  form  a  network  of 
fine  anastomosing  tubules  just  dorsal  to  the  testes  and  within  the  medul- 
lary parenchyma.  The  main  branches  of  the  vasa  efferentia  unite  to 
form  the  larger  vas  deferens.  The  junction  occurs  at  a  point  near  the^ 
median  line  about  one-fourth  the  distance  from  the  anterior  to  the  pos- 
terior end  of  the  segment.  From  this  point  of  juncture  the  vas  deferens 
forms  a  mass  of  coils  extending  to  the  cirrus-pouch.  These  coils  function 
as  a  vesicula  seminalis.  When  the  vas  deferens  enters  the  cirrus-pouch 
it  receives  some  additional -glandular  elements,  the  prostate  cells.  This 
part  of  the  vas  deferens  is  called  the  ductus  ejaculatorius.  The  ductus 
in  this  species  is  very  nearly  straight.  The  cirrus  which  is  a  continua- 
tion of  the  ductus  ejaculatorius  has  a  heavier  musculature  than  is  pres- 
ent in  any  other  part  of  this  canal.  The  cirrus  lies  in  the  pouch  as  an 
almost  straight  organ.  When  protruded  it  measures  0.07-0.08  mm.  long 
by  0.02-0.025  mm.  broad.  The  cirrus-pouch  is  elongated  oval  in  outline. 
It  is  short  and  stout,  being  0.13-0.14  mm.  long  by  0.05-0.06  mm.  broad. 
The  ratio  of  its  length  to  the  proglottid  breadth  is  1 :4  to  1 :3. 

The  vaginal  pore  opens  into  the  genital  sinus  just  dorsal  to  the 
cirrus-pouch  tho  the  greater  part  of  the  initial  part  of  the  vagina  itself 
lies  anterior  to  the  pouch.  A  small  sphincter  vaginae  is  situated  near 
the  opening.  In  its  course  to  the  mid-field  the  vagina  crosses  the  inner 
end  of  the  cirrus-sheath.  When  passing  posteriad  to  enter  the  inter- 
ovarial  space  the  vagina  is  straight,  not  sinuous  or  coiled.  Just  anterior 
to  the  ovary  the  vagina  dilates  slightly  to  form  a  receptaculum  seminis. 
The  relations  of  the  vagina  are  shown  in  figures  88,  90,  and  92.  The 
ovary  is  bi-lobed  .  The  lobes  (Fig.  92)  are  elongated  oval  and  smooth  in 


69]  PROTEOCEPHALIDAE  —  LA  RUE  69 

outline.  The  vitellaria  are  typical  of  the  genus.  The  uterus  is  formed 
of  a  median  ventral-lying  tube  which  extends  nearly  the  full  length  of 
the  proglottid.  From  this  tube,  by  the  process  described  for  Ophiotaenia 
filaroides,  lateral  outpocketings  arise.  These  lateral  pouches  (Fig.  92) 
number  10-14  on  either  side.  As  the  pouches  become  filled  with  eggs 
the  outlines  of  some  of  them  are  nearly  obliterated.  Yet  in  all  but  the 
ripest  proglottids  the  full  number  is  visible.  Sections  through  mature 
and  ripening  proglottids  show  the  pouches  in  all  stages  of  development. 
Sections  also  reveal  the  ventral  diverticula  which  finally  come  to  open 
on  the  surface.  Such  diverticula  were  described  in  an  earlier  paper  (La 
Rue  1909:36)  for  Ophiotaenia  filaroides  and  their  presence  in  other  spe- 
cies was  at  that  time  noted.  In  P.  pinguis  the  number  of  ventral  diver- 
ticula is  greater  than  Gui.  Schneider  (1905)  reported  in  P.  macrocepha- 
lus,  P.  percae,  P.  esocis,  and  P.  ambiguus  but  smaller  than  the  number 
reported  by  the  writer  for  0.  filaroides.  A  drawing  (Fig.  89)  of  a 
frontal  section  just  below  the  cuticula  shows  two  of  these  openings.  In 
such  sections  the  openings,  usually  two  or  three,  and  the  tubes  leading 
up  to  them  could  be  readily  traced.  A  specimen  cleared  in  glycerine 
showed  these  openings  beautifully.  The  actual  count  of  openings  on  25 
consecutive  proglottids  of  this  specimen  is  as  follows :  1,  2,  3,  2,  2,  2,  1, 
1,  2,  3,  2,  2,  2,  2,  2,  3,  2,  4,  3,  2,  4,  2,  3,  3,  3.  The  uterine  eggs  were 
observed  only  in  preserved  condition.  The  outer  mucilaginous  envelope 
could  not  be  seen.    The  embryos  measured  0.016-0.018  mm  . 

Proteocephalus  pinguis  La  Rue  is  a  much  larger  form  than  P.  esocis 
(Gui.  Schneider).  It  has  a  fifth  sucker  which  the  latter  lacks.  It  dif- 
fers from  that  species  in  having  a  larger  head,  larger  suckers,  larger 
proglottids,  more  testes  and  a  relatively  shorter  cirrus-pouch.  P.  pin- 
guis differs  from  P.  pusillus  "Ward,  a  species  occurring  in  the  same  local- 
ity, in  the  possession  of  a  larger  head,  larger  suckers,  larger  proglottids 
which  have  proportions  different  from  those  of  P.  pusillus.  Moreover, 
the  segmentation  is  of  a  different  character  and  the  relative  lengths  of 
the  cirrus-pouches  in  the  two  worms  are  very  different.  P.  pinguis  dif- 
fers very  much  from  P.  exiguus  La  Rue  in  size,  in  the  proportions  of  the 
proglottids,  in  the  size  of  the  head  and  of  the  suckers,  in  the  relative 
length  of  the  cirrus-pouch,  in  the  character  of  the  uterine  pouches  and 
in  the  size  of  the  embryos. 

P.  pinguis  differs  from  P.  macrocephalus  (Creplin)  in  having  a 
fifth  sucker,  in  being  considerably  smaller,  in  the  relative  length  of  the 
cirrus-pouch,  in  the  number  of  testes,  and  in  the  size  of  the  embryos. 
P.  pinguis  is  smaller  than  P.  ambloplitis  (Leidy),  P.  singularis  La  Rue 
and  P.  perplexus  La  Rue.  There  is  also  a  difference  in  the  size  and 
character  of  the  head  and  suckers.     It  is  further  differentiated  from 


70  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [70 

those  species  by  the  presence  of  a  functional  fifth  sucker  which  those 
species  lack.  P.  pinguis  is  quite  different  from  the  European  forms 
which  have  fifth  suckers.  These  are  P.  percae  (O.  F.  Miiller),  P.  fallax 
La  Rue,  P.  duhius  La  Rue,  P.  cernuae  (Gmelin),  P.  longicollis  (Rud.). 
It  differs  from  them  in  size  of  body,  size  of  suckers  and  of  proglottids, 
in  the  relative  length  of  cirrus-pouch,  in  the  size  of  the  embryos  and  in 
the  number  of  testes. 

PROTEOCEPHALUS  FALLAX  La  Rue 
[Figs.  23,  56,  57,  150,  170,  178,  179] 


1884 
1884 
1892 
1892 
1905 
1905 
1911 


Taenia  ocellata  Zschokke  1884:13-14,  in  part ( ?) 

Taenia  longicollis  Zschokke  1884:14-15 

Taenia  filicollis  Kraemer  1892:535-555 

Taenia  ocellata  Kraemer  1892:572-576 

Proteocephalus  ocellata  Nufer  1905  in  part(  ?) 

Proteocephalus  longicollis  Nufer  1905  in  part(  ?) 

Proteocephalus  fallax  La  Rue  1911:475-477 


Specific  Diagnosis :  Characters  of  the  genus.  Cestodes  quite  small. 
Length  up  to  100  mm.  Maximum  breadth  as  much  as  1.20  mm.  Head 
somewhat  globose,  0.150-0.200  mm.  in  breadth  by  0.105-0.150  mm.  thick. 
Head  well  set  off  from  neck.  Suckers  situated  near  broadest  zone  of 
head.  Diameter  of  suckers  0.064-0.085  mm.  Diameter  of  sucker  opening 
about  0.040  mm.  Fifth  sucker  present,  functional,  about  0.058  mm.  in 
diameter.  Neck  narrow,  length  2-6  mm.  First  proglottids  quadrate  or 
longer  than  broad.  Mature  proglottids  quadrate  or  broader  than  long, 
0.34-0.46  mm.  broad  by  0.34  mm.  long.  Ripe  proglottids  longer  than 
broad,  rarely  measuring  as  muclj  as  1.19  mm.  broad  by  1.36  mm.  long. 
Segmentation  indistinct,  margins  smooth. 

Genital  sinus  marginal,  alternating,  situated  near  middle  of  the 
proglottid.  Testes  30-35  in  number,  irregularly  scattered  in  single  layer 
between  vitellaria  and  anterior  to  ovary.  Diameter  of  testes  0.037-0.060 
mm.  Cirrus-pouch  in  mature  proglottids  0.196-0.255  mm.  long,  in  ripe 
proglottids  0.37-0.42  mm.  long.  Ratio  of  length  of  cirrus-pouch  to 
breadth  of  proglottid  1:3-1:2.  Cirrus  and  ductus  ejaculatorius  nearly 
straight.  Length  of  protruded  cirrus  0.026  mm.  Vas  deferens  forming 
a  mass  of  coils  in  middle  of  proglottid. 

Female  organs  typical  of  genus.  Vaginal  opening  anterior  to  cirrus- 
pouch.  Vaginal  sphincter  weak.  Ovary  bilobed,  lobes  solid,  thick,  elon- 
gated ovoidal  in  form.  In  ripe  proglottid  ovary  lies  in  triangular  area 
between  pouches  of  uterus.    Vitellaria  lateral,  sparse.    Uterus  ventral. 


71] 


PROTEOCEPHALIDAE  —  LA  RUE 


71 


in  ripe  proglottids  with  6-8  lateral  pouches  on  either  side.  Uterine  pores 
2-3  in  number.  Outer  membrane  of  uterine  eggs  thin  and  hyaline,  mid- 
dle membrane  granular,  0.036-0.041  mm.,  embryo  0.031-0.0336  mm.  in 
diameter. 

Habitat:  Intestine  of  Cor  eg  onus  fera  (type  host) ;  Lake  Lucerne, 
Switzerland  (type  locality). 

Type:  No.  09.9  in  Professor  H  .B.  "Ward's  collection.  Slides  from 
same  lot  in  Professor  Ward's  collection. 

Zsehokke  (1884)  reported  and  described  in  a  very  general  way  five 
species  of  Proteocephalus.  His  Taenia  salmonis  umhlae  and  Taenia 
torulosa  are  discussed  in  their  proper  places.  The  other  three  species, 
his  Taenia  ocellata,  T.  filicoUis,  and  T.  longicollis  can  best  be  discussed 
together  and  in  this  connection.  The  small  table  shows  the  distribution 
of  the  five  species  in  the  hosts  from  Lake  Geneva  examined  by  him. 


Hosts 

Proteocephalus  Species 

Coregonus 

Taenia 

Taenia 

Taenia 

fera 

longicollis 

ocellata 

torulosa 

Salmo  unibla 

T.  longicol- 
lis 

T.  ocellata 

Taenia  sal- 
monis 

Trutta 

T.  longicol- 

T. ocellata 

umhlae 

variabilis 

lis 

Perca 

T.  ocellata 

Taenia 

fluviatUis 

filicoUis 

Esox  lucius 

T.  ocellata 

Lota  vulgaris 

T.  ocellata 

T.  torulosa 

Alhurnus 

T.  torulosa 

lucidus 

The  Taenia  filicoUis  of  Zsehokke 's  description  (1884:16-17)  is  from 
Perca  fluviatUis  and  may  be  the  same  as  the  specimens  from  the  same 
host  species  received  by  Dr.  H.  B.  "Ward  from  Professor  Parona  which 
La  Rue  (1911)  has  named  P.  dubius.  It  is  certain  that  Zsehokke 's  de- 
scription and  drawings  of  his  specimens  from  this  host  do  not  agree  with 
Schneider's  specimens  of  P.  percae  (Miiller). 

The  specimens  of  T.  ocellata  which  Zsehokke  reported  from  Perca 
fluviatUis  are  probably  only  longer  specimens  of  his  T.  filicoUis.  The 
specimens  reported  by  him  from  Esox  lucius  were  probably  Proteoceph- 
alus esocis  Schneider.  It  is  doubtful  if  Lota  vulgaris  harbors  the  same 
species  of  parasite  as  does  the  Salmonoid  fishes.    Zsehokke 's  report  on 


72  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [72 

this  point  needs  confirmation.  The  Taenia  longicollis  and  T.  ocellata 
from  Coregonus  fera  are  probably  identical.  Zschokke's  description  of 
his  Taenia  longicollis  was  no  doubt  based  on  specimens  from  Coregonus 
fera  for  he  wrote  (p.  14)  :  "Pendant  mes  recherches  je  I'ai  trouve  en 
grande  quantite  dans  les  appendices  pyloriques  et  dans  les  intestines 
greles  de  Coregonus  fera,  et  une  fois,  en  Fevrier,  enkyste  sous  sa  forme 
larvaire  dans  le  foie  de  Salmo  Umbla." 

Fortunately  Dr.  H.  B.  "Ward  has  received  some  specimens  from 
Professor  Zschokke  labelled,  ^'Taenia  ocellata,  Coregonus  fera."  These 
have  been  found  to  be  identical  with  Kraemer's  Taenia  ocellata  from 
Coregonus  fera,  Lake  Lucerne,  and  they  have  proved  to  be  a  new  species 
to  which  the  writer  (La  Rue  1911)  gave  the  name  of  P.  fallax  La  Rue. 
It  is  highly  probable  that  the  specimens  from  Coregonus  fera  identified 
by  Zschokke  (1884)  as  T.  longicollis  belong  to  this  same  species.  The 
T.  longicollis  of  his  description  differs  from  that  of  Kraemer  (1892) 
chiefly  in  length.  The  small  head,  large  cirrus-pouch,  the  proportions  of 
the  proglottids,  position  of  the  genital  pore,  and  the  size  of  the  cirrus- 
pouch  agree  pretty  well  with  the  facts  for  P.  fallax  La  Rue.  Zschokke 
(1884)  presented  no  data  to  show  that  he  had  made  a  comparative  study 
of  the  specimens  collected  by  him  from  Salmo  umhla  and  from  Trutta 
variabilis.  It  is  scarcely  possible  to  determine  where  these  specimens 
belong,  and  speculation  without  some  basis  of  fact  is  worth  but  little. 

Kraemer  (1892)  as  a  result  of  his  study  of  specimens  taken  from 
Coregonus  fera,  Lake  Lucerne,  in  July,  identified  these  specimens  as 
Taenia  filicollis  Rud.  Later  in  the  same  season  (about  Sept.  1)  he  took 
more  specimens  from  Coregonus  fera.  These  he  identified  as  Taenia 
ocellata  on  account  of  their  greater  length.  His  specimens  collected  in 
July  measured  30-60  mm.  Those  taken  by  him  in  September  measured 
as  much  as  100  mm.  Kraemer  now  made  a  careful  comparison  of  these 
two  forms.  He  found  that  they  agreed  in  every  particular  as  to  size  of 
the  head  and  suckers  and  as  to  the  presence  of  a  fifth  sucker.  In  the 
internal  anatomy  of  the  proglottids  and  in  the  histological  structure  of 
the  organs  the  two  forms  also  agreed  perfectly.  Moreover  among  his 
specimens  he  found  gradations  in  length  from  the  shortest  to  the  longest. 
On  these  grounds  Kraemer  concluded  that  Taenia  filicollis  Rud.  and 
Taenia  ocellata  Rud.  were  one  and  the  same;  that  in  fact  the  smaller 
Taenia  filicollis  was  but  an  incompletely  developed  Taenia  ocellata. 
Kraemer  was  correct  in  concluding  that  these  larger  and  smaller  speci- 
mens in  his  possession  belonged  to  the  same  species.  However,  his  con- 
clusion has  nothing  whatsoever  to  do  with  Rudolphi's  Taenia  ocellata  and 
T.  filicollis  for  the  very  good  reason  that  Kraemer's  specimens  belonged 
to  neither  of  Rudolphi  's  species. 


73]  PROTEOCEPHALIDAE  —  LA  RUE  73 

Kraemer  in  his  paper  gave  no  adequate  reasons  for  considering  that 
the  Taenia  filicoUis  and  Taenia  ocellata  of  Rudolphi  were  identical.  His 
specimens  were  neither  from  Rudolphi 's  type  host  nor  from  his  type 
locality  but  from  a  very  different  host  and  locality.  Zschokke  had  ma- 
terial from  Perca  fluviatUis  yet  he  described  what  he  considered  to  be 
Taenia  filicoUis  rather  than  Taenia  ocellata  from  that  host.  Neither 
Zschokke  nor  Kraemer  reported  parasites  from  Gasterosteus  aculeatiis  or 
G.  pungitiiis,  neither  of  which  occur  in  the  waters  from  which  their  fish 
came.  Kraemer 's  specimens  beyond  a  doubt  belong  to  the  species  P. 
fallax  La  Rue.  Riggenbach  (1896)  accepted  Kraemer 's  conclusion  con- 
cerning the  identity  of  Taenia  ocellata  and  Taenia  filicoUis.  He  appar- 
ently made  no  comparative  study  of  the  forms.  From  North  American 
fish  Benedict  (1900)  described  a  species  which  he  considered  identical 
with  Kraemer 's  Taenia  ocellata  and  so  named  it  Proteocephalus  ocellata 
(Rud.).  That  however  was  a  misdetermination,  for  Benedict's  specimens 
have  been  shown  to  belong  to  another  species  (see  P.  exiguus  La  Rue). 
Nufer  (1905)  may  have  been  dealing  in  part  with  this  species  in  his 
statements  regarding  P.  ocellatus  and  P.  longicollis  from  Coregonus  fera. 
La  Rue  (1911 :47 6-477)  described  this  as  a  new  species,  P.  fallax. 

The  material  on  which  this  species  is  based  bears  the  label  "T.  ocel- 
lata, Coregonus  fera."  It  was  secured  from  Prof.  Fritz  Zschokke  by 
Prof.  H.  B.  Ward.  It  now  bears  the  number  09.9  in  Professor  Ward's 
collection.  Some  of  this  material  was  stained  in  haematoxylin  and 
mounted  in  balsam.    Other  specimens  were  cleared  in  glycerine. 

A  comparison  of  these  preparations  with  Kraemer 's  description 
made  evident  the  fact  that  this  species  and  Kraemer 's  Taenia  ocellata 
are  anatomically  identical.  The  fact  that  Kraemer 's  specimens  were 
also  from  Coregonus  fera  and  that  doubtless  they  came  from  the  same 
locality  was  one  of  the  considerations  which  caused  this  identity  to  be 
suspected.  Kraemer 's  descriptions  of  T.  ocellata  will  be  used  as  sources 
of  comparative  data,  all  of  his  data  being  grouped  together. 

The  larger  worms  at  the  writer's  disposal  were  unfortunately 
broken  into  pieces.  One  complete  strobila  measured  27  mm.  long  with 
a  maximum  breadth  of  0.459  mm.  A  small  piece  in  the  same  lot  had  a 
maximum  breadth  of  1.20  mm.  Kraemer  reported  specimens  of  T.  ocel- 
lata 30-60-100  mm.  long.  He  reported  a  breadth  of  0.114  mm.  for  the 
head  and  0.038  mm.  for  suckers.  The  last  measurement  must  be  that 
of  the  sucker  opening  and  not  the  maximum  diameter  of  the  sucker. 


74 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


[74 


Measurements  in  mm.  of  seven  scolices  and  suckers  may  be  seen  in 
the  following  table : 


No. 


Breadth  of  head 


Length 
of  head 


Suckers,  maximum 
diameter 


Fifth 
sucker 


1 
2 
3 

4* 

5 

6 

7 


0.187  mm. 

0.185  mm. 

0.170  mm. 

0.185  mm. 

0.204  mm. 

0.153  X  105  mm.  thick 

0.187  X  0.153  mm.  thick 


short 


0.085  mm. 

0.080  mm. 

0.064  X  0.064  mm. 

0.068  mm. 

0.082  X  0.074  mm. 


not  observed 


0.079  mm. 


0.058  mm. 


There  is  here  some  discrepancy  between  Kraemer's  figures  and 
those  of  the  writer.  This  may  be  due  to  an  error  in  manipulation,  to  the 
fact  that  Kraemer  happened  to  measure  a  very  slender  head,  or  that  he 
happened  to  measure  it  in  the  dorsoventral  (the  shorter)  dimension.  As 
will  be  noted  the  diameter  of  the  sucker  as  given  by  Kraemer  is  about  the 
size  which  the  writer  gives  for  the  sucker  opening.  The  head  (Figs.  23, 
150)  is  somewhat  globular  in  shape.  The  suckers  are  situated  at  or 
just  above  the  broadest  zone  and  they  are  directed  somewhat  anteriad. 
In  the  writer's  material  the  head  is  readily  distinguished  from  the  thin 
neck.  The  neck  is  0.076  mm.  broad  (Kraemer),  or  0.136  mm.  broad  in  the 
writer's  measurements.  Kraemer  states  that  it  is  long  but  he  points  out 
that  in  stained  preparations  part  of  that  which  appeared  to  be  neck 
was  in  reality  made  up  of- young  proglottids.  In  glycerine  mounts  the 
neck  sometimes  appeared  to  the  writer  to  be  as  much  as  6  mm.  long  while 
in  stained  preparation  2.38  mm.  was  the  maximum  length. 

The  first  proglottids  (Fig.  170)  are  about  quadrate  or  they  may  be 
longer  than  broad.  Kraemer  gives  the  following  breadths  for  the  various 
proglottids :  First  long  proglottids  0.228  mm.  broad,  the  quadrate  ante- 
rior proglottids  0.532  mm.,  the  sexually  ripe  mid-proglottids  scarcely 
2.0  mm.,  the  ripest  end-proglottids  as  much  as  2.0  mm.  The  writer's 
measurements  of  similar  proglottids  are  as  follows:  The  first  proglot- 
tids 0.136  mm.  broad  by  0.17-0.30  mm.  long,  quadrate  mature  proglottids 
0.34-0.46  mm.  broad  by  0.34  mm.  long,  the  sexually  ripe  proglottids,  1.19 
mm.  broad  by  1.36  mm.  long  (only  four  proglottids  of  this  size  were 
observed).  The  segmentation  of  the  worm  is  indistinct,  the  margins 
being  quite  smooth  except  for  very  slight  indentations  at  the  junction 
of  the  segments.    The  segments  are  closely  joined  together.    The  genital 

*The  opening  of  the  suckers  in  No.  4  measured  0.042  mm. 


75]  PROTEOCEPHALIDAE  —  LA  RUE  75 

pore  is  not  marked  by  a  papilla.  The  common  genital  sinus  is  marginal, 
irregularly  alternating,  and  situated  near  or  slightly  anterior  to  the 
middle  of  the  segment. 

Kraeraer  worked  out  in  considerable  detail  the  structure  of  the 
cuticula,  the  musculature,  the  parenchyma,  the  nervous  system,  the  ex- 
cretory system,  and  the  histological  structure  of  the  various  internal 
organs.  In  the  main  the  writer  does  not  discuss  these  points,  but  since 
some  of  Kraemer's  conclusions  are  incorrect  the  errors  are  indicated 
when  possible. 

In  preparations  studied  by  the  writer  the  testes  number  30-35  and 
they  measure  0.037  by  0.053  mm.  Kraemer  recorded  27-30  testes  with 
a  diameter  of  0.057  mm.  The  testes  (Figs.  57,  178)  lie  in  a  single  layer 
in  the  field  bounded  by  the  ovary,  the  anterior  margin  of  the  proglottid 
and  the  vitellaria.  The  cirrus-pouch  is  0.196-0.255  mm.  long  in  fully  ma- 
ture proglottids,  where  its  ratio  to  the  proglottid  breadth  is  1 :3-l  :2.  In 
fully  ripe  proglottids  it  is  more  or  less  concealed  by  the  uterine  pouches. 
In  a  few  ripe  proglottids  in  which  it  was  not  contracted  the  cirrus-pouch 
measured  0,370-0.420  mm.  in  length.  Apparently  in  this  species  the 
cirrus-pouch  becomes  larger  after  sexual  maturity  is  attained.  The 
cirrus-pouch  is  slender.  Its  inner  end  is  slightly  curved  upward  toward 
the  dorsal  surface  where  it  is  attached  to  the  walls  of  the  dermo-muscular 
sac  by  strong  muscle  fibers.  This  inner  end  (Figs.  57,  178)  is  usually 
somewhat  swollen  making  a  little  vesicle.  With  this  vesicle  one  or  two 
sinuous  curves  of  the  ductus  ejaculatorius  may  be  seen.  Adjacent  to 
the  vesicle  is  the  constricted  region  of  the  cirrus-pouch.  Here  even  in 
toto  preparations  strong  circular  muscle  fibers  may  be  seen.  In  this  re- 
gion the  ductus  passes  over  into  the  straight  slender  cirrus*  Just  beyond 
this  constricted  region  the  cirrus-pouch  again  dilates  a  little  and  here 
the  circular  muscle  fibers  are  not  prominently  developed.  From  a  cir- 
rus-pouch so  constricted  the  cirrus  itself  is  usually  protruded.  The 
length  of  the  incompletely  protruded  cirrus  is  about  0.026  mm.  Krae- 
mer states  that  the  cirrus  protrudes  2-3  mm.  This  must  be  an  error, 
probably  typographical,  for  a  cirrus  as  long  as  2-3  mm.  would  require 
first  a  very  large  cirrus-pouch  and,  second,  many  coils  of  vas  deferens 
within  the  pouch.  Kraemer's  drawings  and  data  are  not  convincing 
that  this  is  the  case.  His  figure  (Fig.  178)  shows  a  protruded  cirrus 
probably  not  over  0.3  mm.  in  length.  The  vas  deferens  forms  a  mass  of 
coils  at  the  inner  end  of  the  cirrus-pouch.  In  mature  proglottids  this 
knot  lies  in  the  mid-field,  even  reaching  past  the  middle. 

Kraemer  undoubtedly  made  some  errors  in  his  description  of  the 
cirrus  of  this  species.    Benedict  (1900)  pointed  out  these  errors  thus: 


76  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [76 

"Kraemer's  description  is  widely  different.  His  drawing  shows  a  tube  with 
hooks,  representing  the  cirrus,  and  followed  by  a  proglottis-like  arrangement,  and 
then  several  coils  of  the  vas  deferens  within  the  pouch.  The  drawing  has  an  unreal 
appearance  on  the  first  glance.  The  cirrus,  with  its  curved  hooks,  is  imbedded  in 
the  tissue,  which  must  be  torn  through  before  it  could  be  protruded.  The  proglot- 
tis-like appearance  of  a  portion  of  the  cirrus,  as  he  drew  it,  was  undoubtedly  due 
to  the  way  in  which  the  circular  muscles  were  cut  in  sectioning.  He  drew  the 
cavity  of  the  cirrus,  into  which  the  cut  ends  of  the  circular  muscles  projected,  as 
the  external  outline  of  the  cirrus.  The  coils  would  be  necessary,  according  to  his 
theory,  but  are  not  to  be  found.  He  says  that  the  muscles,  which  he  calls  "the 
roots,"  are  for  retracting  the  pouch,  which  is  sometimes  thrust  outwards  for  some 
distance  through  the  opening.  This  protrusion  is  really  due  to  an  evagination  of 
the  free  distal  end  of  the  inner  tube." 

The  female  organs  in  arrangement  are  typical  of  the  genus.  The 
vagina  opens  always  anterior  to  cirrus-pouch.  Near  its  opening  at  a 
distance  of  0.041  mm.  according  to  Kraemer,  it  possesses  a  weak  sphinc- 
ter muscle  0.026  mm.  long  by  0.034  mm.  broad  over  all.  In  its  course  to 
the  interovarial  space  it  crosses  the  bulbous  end  of  the  cirrus-pouch  or 
extends  beyond  the  end  of  the  pouch  before  bending  posteriad.  The 
writer  has  not  noted  in  toto  preparations  the  dilation  in  the  middle  por- 
tion of  the  vagina  as  described  by  Kraemer.  There  are  no  coils  of  the 
vagina  anterior  to  the  ovary.  The  ovary  is  bilobed,  the  lobes  connected 
by  a  slender  mid-piece.  The  lobes  are  solid,  fairly  thick  bodies  of  elon- 
gated ovoidal  shape.  In  mature  proglottids  the  span  of  the  ovary  is 
about  0.350  mm.  In  ripe  proglottids  the  ov^ry  instead  of  becoming  more 
elongated  and  flattened  against  the  posterior  wall  of  the  segment  shrinks 
in  size  and  the  two  lobes  become  quite  closely  pressed  together  in  a  small 
triangular  space  (Fig.  56) -bounded  by  the  posterior  proglottid  margin 
and  by  the  walls  of  thie  uterine  pouches.  This  triangular  space  formed 
by  uterine  pouches  in  the  posterior  end  of  the  proglottid  is  a  very  char- 
acteristic feature  in  this  species.  The  vitellaria  are  sparse  follicular 
glands  in  the  lateral  margins  of  the  proglottid. 

The  organs  of  the  interovarial  space  are  probably  about  as  Benedict 
has  described  them  for  his  species  and  not  entirely  as  Kraemer  has 
described  them.  The  writer  has  not  made  sections  of  this  form  and  has 
been  unable  to  trace  the  connections  of  the  ducts.  Kraemer  established 
the  presence  of  an  ootype,  an  oocapt,  and  a  shellgland.  The  paired 
vitelline  ducts  do  not  originate  as  far  anterior  as  Kraemer  has  figured 
them.  They  probably  unite  to  form  a  single  vitelline  duct  as  in  other 
Proteocephalid  species  where  this  has  been  investigated.  Kraemer  noted 
in  toto  preparations  that  the  vagina  anterior  to  the  ovarj-  was  appar- 
ently divided,  one  branch  going  to  the  ovary.  The  second  tube  is  prob- 
ably the  uterine  passage  which  can  frequently  be  seen  in  toto  prepara- 


77]  PROTEOCEPHALIDAE  —  LA  RUE  77 

tions  and  the  connections  of  which  can  be  worked  out  only  in  recon- 
structions of  sections.  The  uterus  is  a  median  tube  in  mature  proglot- 
tids.  In  ripe  proglottids  (Fig.  56,  179)  there  are  6-8  lateral  pouches 
on  either  side  of  the  median  tube.  These  lie  in  the  ventral  field  and 
come  to  occupy  nearly  the  whole  proglottid.  By  the  pressure  of  the 
densely  packed  uterus  the  vitellaria  are  pushed  farther  toward  the  mar- 
gins; the  testes  are  pushed  close  to  the  dorsal  surface  and  the  shrunken 
ovarie?  are  crowded  into  a  small  triangular  space  in  the  posterior  part 
of  the  segment.  The  ripe  proglottid  is  practically  a  thin  walled  sac 
divided  up  into  compartments  by  the  thin  septa  of  the  uterine  pouches. 

The  eggs  are  discharged  through  2-3  uterine  pores  (Figs.  56,  179) 
or  through  a  rift  down  the  ventral  surface  which  is  caused  by  a  further 
splitting  of  these  uterine  pores.  The  usual  number  of  uterine  pores  is 
2  or  3  but  occasionally  a  proglottid  is  found  with  a  single  pore.  Krae- 
mer  describes  a  single  uterine  pore  in  the  middle  of  the  segment.  The 
eggs  have  a  shell  with  three  membranes,  the  outer  hyaline,  variable  in 
size,  a  middle  heavier  and  more  granular  membrane,  and  an  inner  thin 
membrane  enclosing  the  embryo.  The  embryos  measure  0.031-0.0336  mm. 
in  diameter.  Such  a  wide  variation  is  probably  due  to  the  different 
forms  which  the  embryos  assume.  Some  are  spherical,  others  ovoidal. 
The  middle  membrane  measures  0.036-0.041  mm.  These  measurements 
show  very  well  the  limits  of  variation.  The  outer  membrane  was  not 
measured. 

This  species  is  readily  differentiated  from  P.  percae  by  its  smaller 
head  and  suckers,  more  slender  neck,  and  smaller  proglottids.  It  is  less 
robust  than  P.  percae.  It  differs  from  P.  percae  in  having  a  much 
shorter  cirrus-pouch,  fewer  testes,  a  shorter  ovary,  and  larger  embryos. 
It  is  readily  distinguished  from  P.  torulosus  by  its  smaller  size  and  by 
its  lack  of  a  fifth  sucker.  It  differs  from  the  P.  longicollis  (Rud.)  as  de- 
scribed by  von  Linstow  in  having  a  smaller  head  with  much  smaller 
suckers.  In  the  P.  longicollis  of  von  Linstow  the  testes  are  much  larger 
than  in  P.  fallax  and  they  are  in  two  ( ?)  fields.  The  cirrus  pouch  is 
shorter  in  P.  longicollis.  The  lateral  pouches  of  the  uterus  are  fewer  in 
number  and  the  embryos  are  smaller  than  in  P.  fallax.  P.  fallax  varies 
from  the  P.  cernuae  in  being  more  slender,  with  a  smaller  head  and  less 
muscular  suckers.  The  latter  species  has  broader  but  narrower  pro- 
glottids, a  shorter  cirrus-pouch,  more  numerous  and  much  larger  testes, 
a  longer  span  of  ovary,  and  smaller  embryos.  P.  fallax  differs  from  P. 
flicollis  (Rud.)  in  being  longer,  in  having  a  fifth  sucker,  a  longer  cirrus- 
pouch,  fewer  testes,  and  larger  embryos. 

P.  fallax  much  resembles  P.  exiguus  but  it  differs  from  that  species 
in  having  a  larger  head,  larger  suckers,  a  larger  fifth  sucker,  a  longer 


78  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [78 

neck,  more  numerous  and  larger  proglottids,  fewer  testes,  a  shorter  cir- 
nis-pouch,  and  fewer  uterine  pouches.  The  embryos  are  larger  than  in 
P.  exiguus.  This  species  resembles  P.  pusUlus  Ward.  It  differs  from 
that  species  in  the  character  of  the  segmentation,  in  the  smaller  size  of 
its  suckers,  in  the  position  of  the  genital  aperture,  in  the  smaller  number 
and  size  of  testes,  in  its  longer  cirrus-pouch  and  in  its  fewer  uterine 
pouches.  P.  fallax  resembles  P.  agonis  Barbieri  but  it  differs  from  that 
species  in  being  somewhat  larger,  in  having  a  fifth  sucker,  fewer  testes, 
and  more  uterine  pouches.  P.  fallax  is  larger  than  P.  esocis  (Schneider) 
and  it  has  a  fifth  sucker.  It  moreover  has  smaller  and  less  numerous 
testes  than  P.  esocis.  P.  fallax  most  closely  resembles  P.  duhius  but  it 
differs  from  that  species  in  having  smaller  and  much  fewer  testes,  a 
relatively  shorter  cirrus-pouch,  fewer  uterine  pouches,  a  slightly  smaller 
embryo,  and  a  smaller  middle  egg  membrane.  In  size,  in  size  of  head, 
and  size  of  suckers  they  resemble  each  other  remarkably. 

PROTEOCEPHALUS  NEGLECTUS  La  Rue 
[Figs.  81,  82] 
1911 :    Proteocephalus  neglectus  La  Rue  1911 :477 

Specific  Diagnosis:  Characters  of  genus.  Strobila  incomplete. 
Head,  suckers,  neck  not  observed.  Outline  of  proglottid  rough.  Seg- 
mentation evident.  Genital  pore  on  a  slight  prominence.  Ripe  proglot- 
tids 0.5  mm.  long  by  0.93  mm.  broad  to  0.75  mm.  long  by  1.53  mm.  broad. 
Young  and  mature  proglottids  not  observed. 

Genital  pores  irregularly  alternating,  situated  near  middle  of  lat- 
eral margin  of  proglottid.  Cirrus-pouch  frequently  contracted,  0.185- 
0.265  mm.  long  by  0.080  mm.  broad.  Normal  cirrus-pouch  0.340  mm. 
long.  Cirrus  and  ductus  ejaculatorius  straight.  Vas  deferens  a  mass 
of  coils  in  mid-field  of  proglottid.  Testes  about  75,  spheroidal,  0.042- 
0.064  mm.  in  diameter,  situated  in  a  single  layer  in  space  between  vitel- 
laria.  Vagina  anterior  to  cirrus-pouch,  its  opening  dorsal  to  opening 
of  cirrus-pouch.  Sphincter  vaginae  poorly  developed,  situated  near 
vaginal  opening.  Vagina  crosses  cirrus-pouch  near  middle.  Receptacu- 
lum  seminis  not  observed.  Ovary  bilobed,  lobes  with  smooth  outlines, 
clubshaped.  Organs  of  interovarial  space  not  observed.  Vitellaria  lat- 
eral, follicular.  Vitelline  follicles  small,  compacted.  Uterus  with  7-9 
lateral  pouches  on  either  side.  Uterine  pores  not  seen.  Embryo  0,026- 
0.0265  mm.  in  diameter.    Outer  egg  membrane  0.042-0.047  mm. 

Habitat:  "Forelle"  (Trutta  fario),  type  host,  probably  from  Lake 
Geneva  or  Lake  Lucerne,  Switzerland. 


79]  PROTEOCEPHALIDAE  —  LA  RUE  79 

Type:  Bottled  material  No.  09.10  in  Dr.  H.  B.  Ward's  collection, 
and  slides  of  same. 

Some  pieces  of  strobila  without  heads  were  received  by  Dr.  H.  B. 
Ward  from  Professor  Zschokke  of  Basel.  These  were  labelled  ^'Taenia 
longicollis  Rud.,  aus  Forelle,  Zschokke."  The  bottle  is  now  No.  09.10  in 
Dr.  H.  B.  Ward's  collection.  Four  toto  unflattened  preparations  were 
made  from  this  material.  This  material  formed  the  basis  of  the  prelimi- 
nary report,  La  Rue  (1911 :477). 

Head,  suckers  and  neck  were  not  seen.  A  proglottid  with  eggs  in 
the  uterus  measured  0.5  mm.  long  by  0.93  mm.  broad.  A  ripe  proglottid 
very  full  of  eggs  was  0.75  mm.  long  by  1.53  mm.  broad.  The  outlines  of 
the  segments  are  rough,  with  several  indentations  on  the  margin  of  each. 
Proglottid  limits  are  well  defined  by  the  indentation  at  the  corners  and 
by  the  rounded  corners.    A  slight  prominence  bears  the  genital  pore. 

Genital  pores  alternate  irregularly  in  the  strobila.  They  are  situ- 
ated one  in  the  middle  of  a  lateral  margin  of  each  segment.  This  genital 
pore  leads  into  a  common  atrium  into  which  the  cirrus-pouch  and  vagina 
open.  When  the  cirrus  is  partly  protruded  the  cirrus-pouch  is  con- 
stricted near  its  inner  end  (Fig.  81).  It  is  also  reduced  in  length,  meas- 
ing  in  various  states  of  contraction  0.185-0.190-0.220-0.265  mm.  by  a 
maximum  breadth  of  about  0.080  mm.  In  a  single  proglottid  the  cirrus- 
pouch  was  in  a  normal  condition.  This  is  delineated  (Fig.  82).  This 
cirrus-pouch  measures  0.390  mm.  long.  The  length  of  the  cirrus-pouch 
is  about  14-3^  the  breadth  of  the  proglottid.  Sometimes  the  cirrus  is 
somewhat  protruded  tho  it  has  not  been  seen  completely  protruded.  It 
is  blunt  at  the  tip.  Within  the  cirrus-pouch  the  cirrus  and  ductus 
ejaculatorius  are  straight.  The  one  passes  imperceptibly  into  the  other. 
Coils  of  the  vas  deferens  cannot  be  seen  clearly  but  their  confused  out- 
lines may  be  discerned  in  the  middle  region  of  the  proglottid  left  vacant 
by  the  testes.  The  testes  numbered  75  and  76  respectively  in  two  pro- 
glottids  in  which  they  were  counted.  They  are  spheroidal,  0.042-0.053- 
0.064  mm.  in  diameter.  It  seems  that  they  lie  in  one  plane,  filling  the 
space  between  the  vitellaria  anterior  to  the  ovary. 

The  uterus  (Fig.  81)  is  made  up  of  a  median  ventral  tube  from 
which  7-9  lateral  pouches  arise  on  either  side.  No  uterine  pores  were 
discovered.  The  vitellaria  are  follicular  and  situated  near  the  lateral 
margins  of  the  segment.  The  follicles  are  small  and  quite  compact.  In 
the  present  species  the  vitellaria  are  less  well  developed  than  in  P.  cer- 
nuae.  The  ovary  is  bilobed  and  is  posteriorly  situated.  The  lobes  are 
long,  thick  and  heavy,  and  have  smooth  outlines.  In  ripe  proglottids  the 
span  of  the  ovarian  lobes  may  be  0.60  mm.  The  vagina  (Fig.  82)  tho 
apparently  situated  anterior  to  the  cirrus-pouch  has  its  opening  into 


80     ■  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [80 

the  genital  atrium  directly  dorsal  to  the  cirrus-pouch.  Near  its  opening 
there  is  a  small  sphincter  vaginae  0.016  mm.  long  by  0.010  mm.  thick. 
Normally  in  its  course  the  vagina  is  slightly  bowed  posteriad  across  the 
middle  or  the  inner  end  of  the  cirrus-pouch.  The  course  of  the  vagina 
in  toto  preparations  cannot  be  traced  to  the  interovarial  space.  The 
organs  of  the  interovarial  space  have  not  been  studied.  The  embryos 
measured  0.026-0.0265  mm.  in  diameter,  the  outermost  envelope  0.042- 
0.047  mm.    The  second  membrane  was  not  measured. 

In  some  respects  the  present  species  resembles  P.  fallax  La  Rue. 
The  triangular  shape  of  the  interovarial  space  of  ripe  proglottids  is  much 
alike  in  the  two  species.  So  also  there  is  considerable  resemblance  in 
respect  to  the  uterine  outpocketings.  The  proglottids,  however,  are  as  a 
rule  broader  and  shorter  in  the  present  species.  In  the  number  of  the 
testes  and  their  close  proximity  to  each  other  this  species  is  very  differ- 
ent from  P.  fallax.  The  cirrus-pouch  tho  greatly  resembling  that  of  P. 
fallax  is  relatively  shorter.  The  two  species  are  alike  in  the  character 
of  the  vagina  and  the  sphincter  vaginae.  Ovaries  are  larger  in  the  pres- 
ent species.  The  best  characters  for  distinguishing  the  two  species  are 
the  number  and  size  of  the  testes,  the  relative  size  of  the  cirrus-pouch 
and  the  size  of  the  embryos.  In  some  respects  this  species  resembles  P. 
duhius  La  Rue  but  the  ripe  proglottids  are  a  little  larger  and  somewhat 
thicker  in  the  present  species.  The  cirrus-pouch  in  proglottids  of  the 
same  stage  of  development  may  be  of  nearly  the  same  length.  This 
species  has  more  testes  and  these  are  smaller  than  in  P.  dubius.  The 
chief  difference  lies  in  the  size  of  the  embryos.  This  difference  amounts 
to  0.005-0.007  mm.  The  second  membrane  of  the  eggs  of  P.  dubius  is 
larger  than  the  outer  membrane  of  eggs  of  P.  neglectus. 

The  present  species  differs  from  P.  percae  in  the  length  of  the  cir- 
rus-pouch, in  the  proportions  of  the  proglottids,  number  and  size  of  the 
testes,  and  in  the  size  of  the  embryos.  P.  neglectus  differs  from  P.  lon- 
gicollis  as  described  by  v.  Linstow  in  the  number,  size  and  arrangement 
of  the  testes.  The  arrangement  of  the  testes  of  P.  longicollis  in  two 
fields  sets  that  species  apart  from  all  the  other  species  reported  from 
fish,  if  von  Linstow 's  observations  on  that  point  are  correct.  P.  neg- 
lectus further  differs  from  that  species  in  the  number  of  uterine  pouches 
and  in  the  position  of  the  genital  pore.  P.  neglectus  does  not  agree  well 
with  Zschokke's  (1884:14-16)  description  of  Taenia  longicollis  Rud. 
from  Coregonus  fera.  The  form  which  he  described  is  apparently  iden- 
tical with  the  P.  fallax  La  Rue  which  occurs  in  the  same  host  species. 


81]  PROTEOCEPHALIDAE  —  LA  RUE  81 

PROTEOCEPHALUS  DUBIUS  La  Rue 
[Figs.  20-22,  75-77,  147-149] 


1884 
1884 
1911 


Taenia  filicollis  Zscliokke    1884:16-17 

Taenia  ocellata  Zschokke   1884:13-14  in  part(?) 

Proteocephalus  dubius     La  Rue     1911:476 


Specific  Diagnosis:  Characters  of  the  genus.  Small  cestodes  as 
much  as  40  mm.  long  by  0.80-1.20  mm.  broad.  Segmentation  indistinct. 
Proglottids  about  100  in  number.  Head  small,  variable  in  shape,  well 
set  off  from  neclf.  Length  of  head  0.10-0.14  mm.,  breadth  0.127-0.212 
mm.  Four  sucliers  0.069-0.080  mm.  in  diameter,  cavities  deep,  sucl^er 
outlines  round  or  irregular.  Fifth  sucker  0.026-0.037  mm.  in  diameter. 
Neck  slender  1.8-3.0-3.5  mm.  long  by  0.085-0.140  mm.  broad.  First  pro- 
glottids broader  than  long,  0.20  mm.  broad  by  0.050  mm.  long.  IVIature 
proglottids  quadrate,  broader  than  long,  or  longer  than  broad,  1.02  mm. 
long  by  0.34  mm.  broad  to  0.595  mm.  long  by  0.680  mm.  broad.  Ripe 
proglottids  quadrate,  broader  than  long,  or  usually  longer  than  broad, 
1.19  mm.  long  by  0.680  mm.  broad  to  0.680  mm.  long  by  1.02  mm.  broad. 
End-proglottid,  triangular,  small,  functional. 

Genital  opening  marginal,  near  middle  of  proglottid  or  anterior 
thereto,  irregularly  alternating.  No  genital  papilla.  Testes  in  two  lay- 
ers between  vitellaria,  anterior  to  ovary,  55-60  in  number.  Vas  deferens 
a  mass  of  coils  in  mid-field.  Cirrus-pouch  long,  slender,  0.265-0.370-0.425 
mm.  in  length  by  0.070-0.085  mm.  in  breadth.  Ratio  of  length  of  cirrus- 
pouch  to  proglottid  breadth  5:11-1:2.  Cirrus  slender,  straight,  when 
protruded  short  and  slender.    Ductus  ejaculatorius  straight. 

Vaginal  opening  anterior  to  cirrus-pouch.  Sphincter  vaginae  weak, 
situated  near  vaginal  opening.  Vagina  crosses  cirrus-pouch  near  middle. 
Ovary  bilobed.  Lobes  thick  somewhat  quadrate  in  nearly  mature  pro- 
glottids, smoothly  ovoidal  in  fully  mature  and  past-mature  proglottids. 
Vitellaria  lateral,  sparse,  follicles  small.  Uterus  with  7-10-12-14  lateral 
pouches  on  either  side.  Uterine  pores  1-2  on  ventral  surface.  Eggs  with 
three  membranes,  second  membrane  granular,  thick,  0.042-0.053  mm.  in 
diameter.    Embryos  about  0.032  ram.  in  diameter. 

Habitat :  Intestine  of  Perca  fluviatUis  (type  host) ;  Lake  Geneva 
(type  locality).  Perhaps  this  species  may  occur  also  in  the  same  host 
species  of  other  lakes  and  rivers  in  southern  or  central  Europe. 

Zschokke  (1884:16-17)  described  some  specimens  of  cestode  found 
by  him  in  Perca  fluviatUis  from  Lake  Geneva.  In  some  respects  his 
description  agrees  fairly  well  with  P.  fallax  La  Rue  from  Coregonus 
fera.    However  in  regard  to  the  ovary  there  is  considerable  difference. 


82  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [82 

Moreover  it  scarcely  seems  probable  that  Perca  fluviatilis  and  Corego- 
71US  fcra  normally  harbor  the  same  parasitic  species.  With  this  doubt 
in  mind  some  material  which  Dr.  H.  B.  Ward  has  received  from  Pro- 
fessor Parona  labelled  Taenia  ocellata  from  Perca  fluviatilis  was  care- 
fully investigated.  It  was  found  that  this  material  which  at  first 
glimpse  appeared  to  be  identical  with  P.  fallax  La  Rue  from  Coregonus 
fera  was  after  all  sufficiently  different  to  make  it  a  separate  species 
which  was  described  by  La  Rue  (1911:476)  from  material  on  which  the 
present  description  is  based.  Material  studied  was  alcoholic  material 
in  Professor  Parona 's  collection  labelled  ^'Taenia  ocellata,  Perca  fluvia- 
tilis," locality  not  known,  and  slides  and  alcoholics  of  the  same  in  Prof. 
H.  B.  Ward's  collections. 

These  cestodes  are  of  small  size,  being  short  and  slender.  The 
longest  specimen  in  Professor  Parona 's  material  measured  about  40 
mm.  in  length  by  about  0.80-1.2  mm.  in  maximum  breadth.  In  such  a 
specimen  there  are  about  30  proglottids  with  eggs  in  the  uterus,  12-18 
mature  proglottids,  about  30  which  show  more  or  less  developed  sexual 
organs,  and  very  few  in  which  the  anlagen  of  the  sexual  organs  cannot 
be  seen,  in  all  about  80-90  proglottids.  Segmentation  is  not  very  dis- 
tinct for  the  proglottids  are  joined  by  nearly  their  entire  breadth.  The 
corners  are  slightly  rounded.  Zschokke  stated  that  the  margins  of  the 
proglottids  are  not  rounded  and  their  angles  are  somewhat  rounded; 
that  the  animal  never  presents  a  crenated  appearance  and  that  it  has 
the  appearance  of  a  ribbon.  Zschokke  found  from  60-100  segments  in 
the  strobila.  His  longest  specimen  was  about  4  cm.  long  and  about  1 
mm.  broad. 

The  head  (Figs.  147,  148)  is  of  small  variable  shape,  well  set  off 
from  the  neck.  Frequently  its  anterior  face  is  somewhat  conical,  at 
other  times  flattened.  The  head  is  flattened  dorsoventrally.  Its  length 
varies  from  0.100-0.140  mm.  Its  breadth  is  from  0.127  mm.  in  heads 
turned  slightly  sidewise  to  0.212  mm.  in  slightly  swollen  heads.  At  its 
broadest  part  the  head  bears  four  suckers  which  are  directed  outward 
and  upward.  These  suckers  appear  to  be  variously  placed  on  the  head 
depending  on  the  state  of  contraction.  A  small  fifth  sucker  0.026-0.037 
mm.  in  diameter  is  situated  at  the  apex  of  the  head.  The  four  suckers 
vary  in  size  from  0.069-0.080  mm.  in  diameter.  They  are  usually  nearly 
circular  in  outline  but  may  be  a  little  irregular.  The  cavity  is  deep 
and  irregular  in  shape  or  it  may  be  nearly  round  in  outline.  The  neck 
is  slender,  1.8  mm.  to  3.0-3.5  mm.  in  length  by  0.085-0.140  mm.  in 
minimum  breadth.  The  length  of  the  neck  was  determined  by  measur- 
ing from  the  head  to  the  first  segmentation  visible  in  stained  prepara- 
tions. 


83] 


PROTEOCEPHALIDAE  —  LA  RUE 


83 


The  following  table  shows  dimensions  of  neck,  head  and  suckers  in 
millimeters : 


Neck 

Head 

Suckers               /if^^^ 
Sucker 

Length 

Breadth 

Length 

Breadth 

Diameter 

1.8 
3.4 

? 

? 
3-3.5 

0.130 
0.085 
0.140 
0.130 
0.130 

0.12-0.140 
0.130 
0.100 
? 
0.130 

0.144 
0.12r 
0.159 
0.212 
0.148 

0.080 
0.079 
0.069 
0.069 
0.069 

0.074 
0.074 
0.074 
0.069 
0.069 

0.026 
0.037 
0.037 
0.033 
0.032 

Zschokke's  (1884)  description,  of  the  head,  suckers  and  neck  reads: 

"La  tete  est  petite,  distincte  du  cou,  arrondie  en-avant.  Son  pourtour  est 
garni  de  quatre  ventouses  dont  la  position  relative  est  fort  variable  (see  Figs. 
147-148). 

"Ces  organes  de  fixation  sont  tres  profonds  et  pourvus  d'une  puissante  mus- 
culature. Quelquefois  ils  sont  reunis  sur  une  de  faces,  de  maniere  qu'on  peut 
les  voir  de  face  tous  les  quatre  a  la  fois.  Frequemment  elles  font  saillie  au  dela 
des  pourtour  de  la  tete.  Celle-ci  prend  alors  une  forme  libee  ou  frangee  fort 
variable  selon  le  degre  de  contraction  des  ventouses.  La  tete  presente  ainsi  souvent 
des  preeminences  distribuees  regulierement,  Les  ventouses  peuvent  meme  aflfecter 
la  forme  de  petites  trompes  protractiles. 

"Le  cou  est  long,  filiforme,  epaissi  en-arriere;  il  n'occupe  cependant  jamais 
plus  d'un  cinquieme  ou  d'un  quart  de  la  longueur  totale  de  I'animal,  tandis  que 
chez  le  Taenia  longicollis  il  atteint  facilement  un  tiers." 

There  is  pretty  good  agreement  between  his  description  and  the 
writer's.  In  the  writer's  specimens  the  first  proglottids  are  broader 
than  long,  0.20  mm.  broad  by  0.05  mm.  long.  These  soon  elongate 
somewhat  and  then  the  anlagen  of  the  genital  organs  begin  to  appear. 
The  proglottids  as  they  become  mature  elongate  and  also  increase  in 
width.  They  may  be  much  elongated,  nearly  square  or  broader  than 
long  depending  upon  the  state  of  contraction.  In  the  same  strobila 
the  following  measurements  were  taken  from  anterior  to  posterior  some 
few  proglottids  being  omitted  at  intervals,  measurements  of  length  being 
stated  first  in  each  instance:  1.02  by  0.34  mm.;  0.68  by  0.425  mm.; 
0.595  by  0.425  mm. ;  0.510  by  0.595  mm. ;  0.595  by  0.680  mm.  The  last 
measurement  is  of  the  last  proglottid  without  uterine  eggs.  Ripe  pro- 
glottids may  be  nearly  square,  longer  than  broad  or  broader  than  long. 
They  may  measure  as  much  as  1.19  long  by  0.68  mm.  broad;  0.68  by 

^Head  turned  slightly  sideways. 


84  ILLIXOIS  BIOLOGICAL  MONOGRAPHS  [84 

1.02  mm.;  0.68  by  0.85  mm.  The  end-proglottid  is  triangular  in  out- 
line, the  posterior  end  being  rounded.  It  is  0.425  mm.  broad  by  0.476 
mm.  long.  Apparently  it  is  functional.  No  excretory  pore  could  be 
seen  at  its  posterior  end.  Of  the  proglottids  Zschokke  wrote:  "Les 
premiers  articles  sont  courts,  deux  a  trois  fois  plus  larges  que  longs. 
Les  suivants  deviennent  carres  et  les  derniers  sont  ordinairement  plus 
longs  que  larges.  L'  article  terminal  est  arrondi  en  arriere.  Les  bords 
des  proglottis  ne  sont  pas  bombes;  leurs  angles  sont  tres  peu  emousses. 
L 'animal  ne  presente  ainsi  point  de  crenulure;  il  a  1 'aspect  d'un 
niban." 

The  genital  opening  which  is  situated  near  the  middle  or  slightly 
anterior  to  the  middle  of  the  lateral  margin  of  the  proglottid  alternates 
irregularly.  There  is  no  genital  papilla.  The  testes  (Figs.  75,  77)  are 
nearly  round  or  at  times  slightly  compressed.  They  measure  0.063- 
0.079  mm.  in  diameter  and  they  are  situated  in  two  partial  layers 
which  cover  the  entire  field  betweeq  the  vitellaria  anterior  to  the  ovary. 
Their  number  is  from  55  to  60.  The  vas  deferens  forms  a  small  com- 
pact mass  of  coils  in  the  midfield  of  the  segment.  The  cirrus-pouch 
(Figs.  75,  76,  77)  is  long,  quite  slender,  smooth  in  outline  or  at  times 
somewhat  constricted  near  the  inner  end  by  the  contraction  of  circular 
muscle  fibers.  The  cirrus-pouch  measures  0.370-0.425  mm.  long  in  ripe 
proglottids,  0.265-0.292  mm.  long  in  a  proglottid  with  few  uterine  eggs 
and  0.228-0.265  mm.  in  mature  proglottids  where  the  cirrus  is  being 
protruded.  Its  breadth  is  about  0.070-0.085  nmi.  It  extends  from  5/11 
to  ^  across  the  proglottid  breadth  or  in  some  cases  it  may  extend 
slightly  past  the  middle.  The  cirrus  is  straight  and  slender.  When 
protruded  it  may  extend  aboat  0.1  mm.  past  the  margin  of  the  segment. 
The  protruded  cirrus  is  slender  or,  if  but  slightly  protruded,  conical. 
The  ductus  ejaculatorius  is  straight  and  of  the  same  size  as  the  cirrus 
of  which  it  really  forms  a  part. 

The  vagina  opens  into  the  conmion  genital  sinus  apparently  ante- 
rior to  the  cirrus-pouch.  Near  its  opening  it  possesses  a  small  sphincter 
vaginae.  The  course  of  the  vagina  is  slightly  anteriad  then  posteriad 
and  mesad  in  a  long  arc  which  crosses  the  cirrus-pouch  near  the  middle 
of  its  dorsal  side.  Its  entire  course  into  the  interovarial  space  has  not 
been  traced.  A  receptaculum  seminis  has  not  been  observed.  The 
ovary  is  posterior  and  is  bilobed,  but  in  nearly  mature  proglottids 
the  mid-piece  may  not  be  visible.  The  lobes  (Fig.  75)  may  appear 
as  more  or  less  irregular  quadrate  bodies  in  the  posterior  comers  of  the 
segment.  In  such  a  condition  they  resemble  the  ovaries  which  Zschokke 
delineated.  His  figure  is  reproduced  (Fig.  149).  In  more  mature  and 
ripe  proglottids  the  lobes  of  the  ovary  are  smoother  in  contour  (Figs. 


85]  PROTEOCEPHALIDAE  —  LA  RUE  85 

76,  77).  The  ovarian  lobes  in  ripe  proglottids  have  a  span  of  0.425 
mm.  The  lobes  are  heavier  and  thicker  than  in  P.  fallax.  The  organs 
of  the  interovarial  space  have  not  been  observed.  In  the  lateral  fields 
of  the  proglottid  lie  the  vitellaria,  made  up  of  small  loose  follicles. 
Well-filled  uteri  (Fig,  76)  are  made  up  of  a  median  tube  and  7-8-10-12 
or  even  14  irregular  lateral  pouches  on  either  side.  The  more  common 
number  of  pouches  is  8  or  9.  One  large  ventral  uterine  pore  occurs 
quite  regularly  near  the  middle  of  the  proglottid  and  a  smaller  pore 
can  sometimes  be  found  farther  anteriad.  Uterine  eggs  have  three 
membranes.  An  outermost  hyaline  membrane  is  variable  in  size  and 
was  not  measured.  The  second  membrane  is  thicker,  is  granular  and 
more  nearly  spherical.  It  has  a  diameter  of  0.042-0.048-0.053  mm.  tho 
its  more  common  dimension  is  0.048  mm.  An  innermost  thin  membrane 
immediately  invests  the  embryo.  The  embryo  has  a  diameter  of  about 
0.032  mm.  but  when  elongated  it  may  measure  0.026  by  0.037  mm.  or 
even  0.032  by  0.037  mm.  The  average  measurement  is  about  0.032  mm. 
Zschokke's  description  of  the  genital  organs  reads  thus: 

"Les  orifices  genitaux  se  trouvent  au  fond  d'un  bourrelet  circulaire,  situe  au 
milieu  ou  un  peu  au-dessus  des  bords  lateraux.  L'alternance  de  la  position  a 
gauche  ou  a  droite  est  irreguliere.  Les  cirrhes  sont  courts,  coniques,  leurs  poches, 
etroites.  Rudolphi  parle  de  ovaria  quadrangularia ;  en  effet  le  parenchyme  est 
rempli  de  vesicules,  qui,  dans  les  articles  jeunes,  ont  une  forme  ronde  ou  ovalaire 
et  deviennent,  dans  les  proglottis  plus  murs,  carrees  et  anguleuses.  En  realite  ces 
vesicules  sont  a  I'etat  normal  pyriformes ;  leur  aspect  carre  est  le  resultat  de  leur 
pression  reciproque.  Du  reste  ce  ne  sont  pas  des  ovaries,  ce  sont  des  testicules 
dont  chacun  presente  un  petit  canal  deferent.  Tous  ces  canalicules  se  reunissent 
dans  un  canal  excreteur  commun. 

"Deja  Dujardin,  en  citant  la  description  de  Rudolphi,  met,  derriere  "ovaires 
opaques,  egalement  carres",  un  point  d'interrogation  et  Diesing  n'  en  parle  plus. 

"Le  vagin  est  court,  il  aboutit  a  une  poche  seminale  assez  spacieuse.  L'uterus 
sous  forme  d'un  tube  large  suit  la  ligne  mediane  du  proglottis  et  se  rend  en 
decrivant  quelques  faibles  lacets  vers  le  bord  posterieur  de  I'article.  La  il  decrit 
un  noeud  de  lacets  et  se  divise  enfin  en  deux  branches,  qui,  de  leur  cote,  con- 
stituent a  droite  et  a  gauche  une  masse  d'ovaires  lobes  ou  contournes.  Entre  les 
ovaires,  touchant  le  bord  posterieur  du  proglottis,  se  trouve  le  glande  vitellogene. 
Des  amas  glandulaires,  peut-etre  la  continuation  directe  des  vitellogenes,  se  trou- 
vent le  long  des  bords  lateraux." 

This  species  is  much  smaller  than  P.  percae  (Miiller).  It  is  most 
closely  related  to  P.  fallax  from  Coregonus  fera.  It  differs  from  that 
species  in  having  regularly  a  larger  number  of  testes  (about  twice  as 
many  as  are  found  in  P.  fallax)  and  these  are  also  larger.  In  P.  fallax 
the  testes  are  in  one  layer,  in  this  species  in  two  partial  layers.  The 
cirrus-pouch  is  usually  longer  than  in  P.  fallax  and  it  is  relatively 


86  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [86 

longer  in  proportion  to  the  proglottid  width.  The  uterine  pouches  may 
be  more  numerous  and  the  embryos  a  trifle  larger  than  in  P.  fallax.  The 
second  egg  membrane  is  larger  in  this  species.  In  other  respects 
the  species  are  much  alike.  Staining  and  careful  comparisons  are  nec- 
essary to  distinguish  them.  P.  duhius  may  be  differentiated  from  P. 
neglectus  by  the  smaller  size  of  its  proglottids  and  by  the  larger  size  of 
its  embryos.  The  proportions  of  the  cirrus-pouch  are  also  very  differ- 
ent. P.  duhius  is  much  unlike  P.  percae  and  P.  cernuae  in  size,  in  size 
of  suckers ;  in  number  and  size  and  arrangement  of  testes,  and  in  length 
of  cirrus-pouch.  This  species  differs  from  P.  esocis,  P.  agonis  and  P. 
filicollis  in  size  and  in  the  presence  of  a  fifth  sucker.  There  is  also  con- 
siderable difference  in  the  length  of  the  cirrus-pouch. 


PROTEOCEPHALUS  CERNUAE  (Gmelin)  La  Rue 
[Figs.  5,  6,  66-68] 


1790: 

Taenia  cernuae 

Gmelin 

1790:3079,  No.  79 

1803: 

Halysis  cernuae 

Zeder 

1803 :376 

1810: 

Taenia  ocellata 

Rudolphi 

1810 :108,  in  part 

<?)1835: 

Taenia  ocellata 

VonSiebold 

1835 :83 

(?)1897: 

Taenia  ocellata 

Von  Ratz 

1897:155,162 

(?)1897: 

Taenia  filicollis 

Von  Ratz 

1897:155,  162 

1898: 

Ichthyotaenia  ocellata 

Miihling 

1898:37 

1911: 

Proteocephalus  cernuae 

La  Rue 

1911 :475-476 

Specific  Diagnosis:  Characters  of  genus.  Strobila  short,  robust. 
Observed  length  as  much  as  40  mm.  Maximum  breadth  1.50  mm.  Num- 
ber of  proglottids  50-60.  Segmentation  not  plain.  Head  not  well  set 
off  from  neck,  flattened  dorsoventrally,  and  with  a  flat  anterior  face. 
Breadth  of  head  0.291-0.316  mm.,  length  about  1.10  mm.  Suckers  not 
prominent,  directed  anteriorly,  placed  above  broadest  zone  of  head. 
Suckers  almost  globular,  heavily  muscled,  small,  with  deep  cavities. 
Diameter  of  suckers  0.064-0.090  mm.  Fifth  sucker  present,  0.024  mm. 
in  diameter.  Neck  0.3-0.39  mm.  broad,  1.7-2.0  mm.  long.  First  pro- 
glottids much  broader  than  long,  0.425  mm.  broad  by  0.085  mm.  long. 
Mature  and  ripe  proglottids  broader  than  long.  Length  exceeds 
breadth  only  in  old  ripe  proglottids.  Dimensions  of  mature  proglottids 
about  0.476  mm.  long  by  0.816  mm.  broad,  ripe  proglottids  1.309-1.51 
mm.  broad  by  0.68-0.85  mm.  long.  End-proglottid  present  and  func- 
tional. 

Common  genital  sinus  situated  near  middle  of  lateral  margin  of 
segment,  irregularly  alternating.     Cirrus-pouch  0.185-0.228  mm.  long. 


87] 


PROTEOCEPHALIDAE  —  LA  RUE 


87 


extending  barely  through  vitellaria.  Ratio  of  length  of  cirrus-pouch  to 
breadth  of  proglottid  2:9  in  mature,  1:5,  1:6,  1:7  in  ripe  proglottids. 
Ductus  ejaculatorius  in  1-3  coils.  Vas  deferens  forming  a  narrow  mass 
extending  to  middle  of  proglottid.  Testes  about  70  in  number,  in  one 
layer,  occupying  entire  field  between  vitellaria  and  anterior  to  ovary. 
Diameter  of  testes  0.085-0.130  mm.  Vagina  anterior  to  cirrus-pouch, 
never  crossing  the  latter.  Sphincter  vaginae  and  reeeptaculum  seminis 
not  seen.  Lobes  of  ovary  thick,  heavy,  and  long,  length  of  pair  0.510- 
0.918  mm.  Vitellaria  coarsely  follicular,  voluminous,  situated  farther 
from  margin  of  proglottid  than  in  most  species.  Uterus  in  ripe  pro- 
glottids with  6-8-9-12  lateral  pouches  on  either  side.  Embryo  with 
three  membranes,  second  membrane  0.037-0.04  mm.  in  diameter,  embryo 
ovoidal,  0.0212-0.0265  mm.  in  length. 

Habitat:    In  intestine  of  Acerina  cernua  (Linn.). 


Host 

Locality 

Collector 

Authority 

Acerina  cernua  (Linn.) 

Pallas 

Gmelin  1790:3079.* 

(?)       " 

Prussia 

Von  Siebold 

Von  Siebold  1835 :83 

(?)       " 

ii 

Liihe 

Muhling  1898 :37 

(?)       " 

Lake 
Balaton 

Von  Ratz 

Von  Ratz  1897 :162 

(?)       "       schraetzer 

<  i 

<< 

Von  Ratz  1897  :162 

' '       cernua 

Konigsberg 

Braun 

Tia  Rue  (the  present 
paper) 

Gmelin  (1790:3079)  refers  to  this  species  in  these  words:  "Taenia 
cernuae.  79 — T.  Pallas  elench.  zooph.  p.  414.  Habitat  in  percae  cer* 
nuae  intestinis,  vix  propria  species."  The  writer  has  not  been  able  to 
secure  the  paper  cited  by  Gmelin  so  he  can  not  judge  as  to  his  descrip- 
tion if,  indeed,  he  gives  one.  Zeder  (1803 :376)  adds  nothing  descriptive 
to  Gmelin 's  data.  He  remarks,  "1st  schwerlich  eine  eigene  Art,  und 
gehort  zu  nro.  42.  (Halysis  percae  mihi)."  Evidently  the  confusion  of 
this  species  with  Proteocephalus  percae  begins  here.  Rudolphi  (1810: 
108)  considers  T.  cernuae  a  synonym  of  Taenia  ocellata  Rud.  (Vide 
infra  for  quotation). 

Von  Siebold  (1835:83)  found  Taenia  ocellata  in  Acerina  cernua  in 
1834.  This  reference  is  from  Muhling  (1898:37).  The  writer  has  not 
seen  von  Siebold 's  paper.    Miihling  also  states  that  Liihe  in  1893  found 

♦Gmelin  gave  this  on  the  authority  of  Pallas  1776:414.  The  writer  has  not 
access  to  this  work  of  Pallas. 


88  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [88 

this  species  in  Acerina  cernua  but  the  writer  has  not  been  able  to  find 
any  reference  to  this  catch  in  any  of  Liihe's  papers.  Von  Ratz  (1897: 
162)  names  Ichthyotaenia  ocellata  (Rud.)  and  /.  filicollis  (Rud.)  as 
parasites  of  Acerina  cernua  Linn,  and  I  filicollis  (Rud.)  as  a  parasite 
of  A.  schraetzer  Cuv.  in  Lake  Balaton.  Since  he  gives  no  description  a 
determination  of  the  systematic  position  of  his  specimens  cannot  be 
made.  They  may  belong  to  P.  cernuae  and  they  probably  do  not  be- 
long to  P.  percae  (Miiller).  Miihling  (1898:17)  stated  that  Ichthyo- 
taenia ocellata  had  been  found  in  East  Prussia  by  Liihe.  La  Rue  (1911: 
475-476)  described  briefly  P.  cernuae  from  specimens  used  in  the  pres- 
ent description. 

Von  Siebold's  and  Liihe's  specimens  were  collected  in  Prussia,  the 
same  general  locality  as  those  of  Braun  which  the  writer  has  had  for 
study.  For  this  reason  and  for  the  reason  that  they  came  from  the 
same  host  species  it  is  deemed  probable  that  they  belong  to  the  same 
species.  The  specimens  collected  by  von  Ratz  were  taken  from  hosts 
occurring  in  a  different  drainage  system  and  may  or  may  not  belong  to 
P.  cernuae.  They  probably  do  not  belong  to  P.  percae  which  comes 
from  the  same  general  region,  as  does  P.  cernuae  and  which  seems  to  be 
limited  to  Perca  fiuviatilis  as  a  host.  It  is  possible,  of  course,  that  they 
represent  a  new  species.  With  these  possibilities  in  mind  von  Ratz's 
specimens  should  be  compared  with  parasites  from  Perca  fiuviatilis  and 
Acerina  cernua  taken  in  Prussia  or  Finland. 

The  description  of  this  species  is  based  on  five  specimens  which 
Prof.  H.  B.  Ward  secured  from  Prof.  Max.  Braun.  The  material  bears 
the  label  '  ^  Proteocephalus  ocellata  Rud.  int.  Acerina  cernua  L.  Konigs- 
berg,  Pr,  8/93."  These  specimens  bearing  the  number  09.26  and  slides 
of  the  same  are  now  to  be  found  in  Professor  Ward's  collection. 

In  general  appearance  these  worms  are  short  and  robust.  Four  of 
them  are  very  short,  10.0-19.5-22.0  mm.  The  fifth  measured  about  40 
mm.  Yet  each  strobila  has  proglottids  with  ripe  eggs.  A  maximum 
breadth  of  1.50  mm.  was  observed  in  the  longest  specimen  and  a  breadth 
of  1.309  mm.  in  a  shorter  one.  The  head  (Figs.  5,  6)  is  not  well  set 
off  from  the  neck.  Indeed,  it  is  continuous  with  the  neck  which  in  all 
five  specimens  is  broader  than  the  head.  The  anterior  face  of  the  head 
is  flattened  and  the  suckers  (Figs.  5,  6)  are  set  at  the  margin  of  the 
flattened  area.  They  are  small  and  not  prominent  and  are  directed 
anteriad.  No  furrows  or  ridges  mark  the  head.  A  fifth  sucker  is  very 
faintly  visible.  The  head  is  0.291-0,316  mm.  broad  at  a  point  a  little 
posterior  to  the  suckers.  It  is  flattened  dorsoventrally.  Its  length 
which  cannot  be  determined  with  any  accuracy  is  about  0.100  mm. 


PROTEOCEPHALIDAE  —  LA  RUE 


89 


The  suckers  are  heavily  muscled,  deeply  concave,  small  and  nearly 
globular  in  shape.  Six  suckers,  four  from  one  head  and  two  from  a 
second,  gave  the  following  measurements: 

Suckers  of  Proteocephalus  cernuae 


Length 

Breadth 

Diameter  of  opening 

0.07  mm. 
0.07  mm. 
0.079  mm. 
0.064  mm. 

0.079  mm. 
0.070  mm. 
0.064  mm. 
0.079  mm. 
0.085  mm. 
0.090  mm. 

0.042  mm.  approximately 
0.042  mm. 
0.037  mm. 

0.090  mm. 
0.090  mm. 

0.037  mm. 
0.037  mm. 

The  range  of  length  and  breadth  of  the  suckers  is  about  0.064-0.090 
nun.  A  sucker  in  transverse  section  measured  0.074  mm.  in  diameter. 
The  sucker  opening  it  will  be  observed  is  very  small,  0.037-0.042  mm. 
The  fifth  sucker  which  seems  to  be  a  true  sucker  is  about  0.024  mm.  in 
diameter.  Its  cavity  is  very  shallow.  Because  of  the  scarcity  of  mate- 
rial no  sections  were  made  and  its  true  structure  could  not  be  finally 
determined.  The  neck  in  all  five  specimens  is  broad  and  thick,  narrow- 
est just  posterior  to  the  head.  Here  it  measures  about  0.3-0.39  mm.  Its 
length  varies  from  1.36-1.7-2.0  mm.  from  the  tip  of  the  head  to  the  first 
traces  of  segmentation. 

The  total  number  of  proglottids  in  one  of  the  shorter  strobilas  was 
52,  in  the  longest,  66.  The  segmentation  is  not  plain.  The  edges  of  the 
chain  are  quite  smooth  except  for  now  and  then  small  folds  which  rarely 
occur  at  the  junction  of  two  proglottids.  The  angles  between  proglot- 
tids are  scarcely  noticeable.  No  longitudinal  folds  or  furrows  were 
seen.  The  first  proglottids  are  much  broader  than  long.  In  one  speci- 
men they  measured  0.425  mm,  broad  by  0.085  mm.  long.  As  the  pro- 
glottids become  older  they  increase  both  in  length  and  breadth.  In  all 
but  a  very  few  ripe  proglottids  the  breadth  exceeds  the  length  or  the 
segments  are  nearly  quadrate.  Mature  proglottids  are  broader  than 
long,  about  0.476  mm.  long  by  0.816  mm.  broad.  Ripe  proglottids  meas- 
ure as  much  as  1.309  mm.  broad  by  0.68  mm.  long  or  even  1.51  mm. 
broad  by  0.85  mm.  long.  One  ripe  proglottid  which  had  discharged 
some  of  its  eggs  measured  1.19  mm.  long  by  0.915  mm.  broad.  A  true 
functional  end-proglottid,  pointed  at  the  posterior  end,  was  1.19  mm. 
broad  at  the  anterior  end  and  1.105  mm.  long.  The  measurements  given 
are  representative  hence  it  will  be  noted  that  in  general  the  breadth 
of  the  proglottids  exceeds  the  length.    Only  rarely  and  then  only  in  old 


90  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [90 

ripe  proglottids  does  the  length  exceed  the  breadth.  The  nervous  and 
excretory  systems,  and  the  musculature  have  not  been  studied. 

The  common  genital  sinus  (Figs.  66,  67)  is  situated  very  near  the 
middle  of  the  lateral  margin  of  the  segment  or  in  ripe  proglottids 
slightly  posterior  to  the  middle.  It,  alternates  irregularly.  There  is  no 
genital  papilla.  In  a  worm  of  52  proglottids  the  anlagen  of  the  sexual 
organs  appear  in  the  10th-12th  segment  as  two  darkly  staining  masses 
the  one  representing  the  cirrus  and  vagina  and  the  other  the  ovaries. 
In  the  20th  segment  the  anlagen  of  the  testes  appear.  Beginning  with 
the  24th  segment  the  sexual  organs  are  mature  while  with  segment  40 
a  number  of  eggs  begin  tq  appear  in  the  uterus. 

The  cirrus-pouch  (Figs.  66,  67)  in  mature  proglottids  is  about 
0.221  mm.  long  while  in  ripe  proglottids  its  length  measures  0.185-0.212- 
0.228  mm.  It  is  an  elongated  ovoid  in  shape  being  broadest  near  the 
inner  end.  In  mature  proglottids  it  extends  just  a  short  distance 
through  the  vitellaria.  In  ripe  proglottids  it  may  barely  reach  through 
the  viteUaria.  The  ratio  of  the  length  of  the  cirrus-pouch  to  the 
breadth  of  the  proglottid  varies  from  2 :9  in  mature  to  1 :5,  1 :6,  1 :7  in 
ripe  proglottids.  The  protruded  cirrus  has  not  been  seen.  "Within  the 
cirrus-pouch  the  distal  portion  of  the  cirrus  is  slender  and  straight. 
The  ductus  ejaculatorius  is  thrown  into  1-3  coils.  The  vas  deferens  in 
scarcely  mature  proglottids  is  thrown  into  coils  which  extend  in  a 
straight  narrow  mass  nearly  to  the  middle  of  the  proglottid,  never  past 
the  middle.  In  ripe  proglottids  coils  of  the  vas  deferens  are  massed 
at  the  inner  end  of  the  cirrus-pouch  and  there  is  no  large  mass  of  coils 
in  the  middle  of  the  segmeijt.  In  P.  fallax  La  Rue  the  coils  of  the  vas 
deferens  form  a  dense  mass  in  the  mid-field  of  the  segment.  In  P.  neg- 
lectus  La  Rue  the  large  mass  of  coils  of  vas  deferens  lies  in  the  middle 
of  the  proglottid.  The  testes  (Fig.  67)  about  70  in  number,  apparently 
lie  in  one  layer.  They  are  irregularly  scattered  between  the  vitellaria 
anterior  to  the  ovary.  They  measure  0.085-0.130  mm.  in  diameter.  They 
are  thus  more  numerous  and  larger  than  in  P.  fallax  and  P.  duhius, 
somewhat  more  numerous  and  larger  than  in  P.  percae  and  much  larger 
than  in  P.  neglectus. 

The  vagina  (Figs.  66,  67)  always  opens  anterior  to  the  cirrus- 
pouch  which  it  never  crosses  in  its  course  to  the  interovarial  space.  It 
passes  to  the  middle  of  the  proglottid  in  a  smooth,  gentle  curve,  then 
it  bends  posteriad  toward  the  interovarial  space.  There  are  no  coils  of 
vagina  anterior  to  the  ovary  but  at  that  place  it  may  be  slightly  sinu- 
ous. This  straight  or  nearly  straight  condition  of  the  vagina  is  evi- 
dence that  the  worm  is  in  a  normal  state  of  contraction  for  usually 
when  a  proglottid  is  unduly  contracted  the  vagina  is  very  sinuous.    In 


91]  PROTEOCEPHALIDAE  —  LA  RUE  91 

an  elongated  proglottid  of  the  same  worm  the  vagina  may  be  straight 
or  nearly  so.  Thus  far  a  sphincter  vaginae  has  not  been  demonstrated. 
The  lumen  of  the  vagina  from  its  opening  to  the  ovary  is  almost  con- 
stant in  diameter.  No  receptaculum  seminis  has  been  demonstrated. 
The  ovary  is  bilobed  as  in  other  Proteocephalids.  The  lobes  are  thick, 
heavy  and  long.  In  all  dimensions  they  are  much  larger  than  in  P. 
fallax  or  P.  duhius.  They  also  have  a  different  outline.  In  mature 
proglottids  they  measure  0.510  mm.  long  while  in  ripe  proglottids  their 
span  may  be  as  much  as  0.918  mm.  This  is  much  greater  than  in  P. 
fallax  or  P.  duhius  and  more  nearly  like  the  condition  in  P.  percae.  The 
organs  within  the  interovarial  space  have  not  been  investigated. 

The  vitellaria  (Fig.  67)  are  coarsely  follicular,  lateral  masses.  The 
follicles  are  much  coarser  and  more  numerous  than  in  P.  duhius  or  P. 
fallax.  The  vitellaria  are  situated  at  a  distance  of  0.100  mm.  from  the 
margin  in  mature  and  ripe  proglottids.  This  is  considerably  farther 
than  in  P.  fallax  and  P.  duhius  and  greater  than  in  P.  percae.  The 
uterus,  a  median  tube  in  mature  proglottids,  is  augmented  in  ripe  pro- 
glottids by  6-8-9-12  lateral  outpocketings  on  either  side.  The  pouches 
(Figs.  66,  68)  are  long,  reaching  to  the  vitellaria,  and  are  voluminous. 
Sometimes  the  larger  pouches  are  apparently  subdivided  by  short  septa. 
For  this  reason  the  appearance  of  the  ripe  proglottid  in  toto  prepara- 
tions differs  considerably  from  that  of  P.  percae  or  P.  duhius.  The 
embryo  is  surrounded  by  three  membranes,  an  outer  thin  and  hyaline, 
a  middle  thick  and  somewhat  granular,  and  an  inner  very  thin  mem- 
brane immediately  investing  the  embryo.  The  embryo  varies  from 
ovoidal  to  spherical  in  shape.  The  measurements  of  four  embryos  may 
be  taken  as  fairly  representative  of  the  range  of  size — 0.0265  by  0.024 
mm.,  0.0212  by  0.0212  mm.,  0.0265  by  0.0212  mm.,  0.024  by  0.0212  mm. 
The  second  membrane  measures  0.037-0.040  mm.  in  diameter.  The 
outer  membrane  is  very  variable  in  size  and  in  the  material  studied 
almost  impossible  to  measure  on  account  of  its  collapsed  condition. 

This  species  somewhat  resembling  P.  percae  and  P.  torulosus  in  its 
outward  appearance  is  distinguished  from  the  latter  by  its  fifth  sucker, 
its  smaller  suckers,  its  shorter  length,  its  much  more  numerous  lateral 
uterine  pouches,  its  fewer  testes,  and  by  the  different  arrangement  of 
testes.  From  P.  percae  it  is  distinguished  by  its  more  numerous  uterine 
pouches,  its  more  numerous  and  larger  testes,  its  much  shorter  cirrus- 
pouch,  and  by  the  different  relations  of  cirrus-pouch  and  vagina.  From 
P.  duhius  and  P.  fallax  it  is  distinguished  by  its  larger  head,  its  pro- 
glottids of  greatly  different  proportions,  its  more  numerous  and  larger 
testes,  by  the  greatly  different  cirrus  and  cirrus-pouch,  by  the  different 
relations  of  the  cirrus-pouch  and  vagina,  by  the  differently  placed  coils 


92  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [92 

of  vas  deferens,  by  the  difference  in  the  size  of  ovaries,  and  by  the 
shape  of  the  ovarial  space  in  the  ripe  proglottid.  From  P.  longicoUis, 
as  described  by  von  Linstow,  P.  cernuae  is  distinguished  by  the  much 
shorter  neck,  the  different  position  of  the  genital  opening,  by  the  num- 
ber of  testes,  by  the  position  and  shape  of  the  mass  of  coils  of  the  vas 
deferens,  by  the  number  of  uterine  pouches,  and  by  the  size  of  the  eggs. 
No  American  species  yet  discovered  resembles  it  closely. 

This  species  is  closely  allied  to  the  form  which  Zschokke  designated 
as  P.  longicollis  (Rud.)  but  which  has  been  determined  to  be  a  separate 
species,  P.  neglectus.  Unfortunately  the  heads  of  P.  cernuae  and  P. 
neglectus  cannot  be  compared  due  to  lack  of  heads  in  the  latter  mate- 
rial. The  shape  and  size  of  the  proglottids  may  be  about  the  same.  In 
P.  neglectus  the  segmentation  is  evident  but  very  indistinct  in  this  spe- 
cies. The  minimum  length  of  the  cirrus-pouch  of  P.  neglectus  is  greater 
than  the  maximum  of  the  same  organ  in  P.  cernuae.  The  ductus  ejacu- 
latorius  in  P.  neglectus  is  straight  but  it  has  from  two  to  three  coils  in 
this  species.  In  ripe  proglottids  of  P.  neglectus  the  ratio  of  the  length 
of  the  cirrus-pouch  to  proglottid  breadth  is  1 :3  to  2 :9  but  in  P.  cernuae 
it  is  1 :5,  1 :6,  1 :7. 

The  coils  of  the  vas  deferens  are  massed  close  to  the  cirrus  pouch 
in  P.  cernuae  but  they  are  mostly  median  in  P.  neglectus.  The  testes  of 
P.  cernuae  are  nearly  double  the  size  of  those  in  P.  neglectus,  so  also 
the  length  of  the  ovary  in  ripe  proglottids  is  much  greater  in  the  former 
than  in  the  latter.  The  extent  and  compactness  of  the  vitellaria  and 
their  relation  to  the  margin  of  the  proglottid  are  different  in  the  two 
species.  There  may  be  more  uterine  outpocketings  in  P.  cernuae  than 
in  P.  neglectus.  The  embryo  measures  0.026-0.0265  mm.  in  P.  neglectus 
and  about  0.021  mm.  in  spherical  embryos  of  P.  cernuae.  A  maximum 
measurement  of  0.0265  mm.  may  occur  in  the  latter  species  but  only  in 
elongated  embryos.  These  species  which  seem  much  alike  at  first  glance 
are  thus  after  all  quite  different. 


93] 


PROTEOCEPHALIDAE  —  LA  RUE 


93 


PROTEOCEPHALUS  PERCAE  (Miiller) 


[Figs.  8,  9,  69-74,  120-122,  172, 

173] 

1780 

Taenia  percae 

Miiller 

1780:152-155,179 

1781 

Taenia  cystica 

Pallas 

1781 :101 

1786 

Taenia  percae 

Batsch 

1786:234-235 

1788 

Taenia  percae 

Miiller 

1788 :5 

1788 

Taenia  percae 

Schrank 

1788 :48 

1790 

Taenia  percae 

Gmelin 

1790:3079 

1802 

Taenia  ocellata 

Rudolphi 

1802a  :112 

1803 

Halysis  percae 

Zeder 

1803 :355 

1810 

Taenia  ocellata 

Rudolphi 

1810:108. 

1819 

Taenia  ocellata 

Rudolphi 

1819 :149 

1845 

Taenia  ocellata 

Dujardin 

1845 :583 

1850 

Taenia  ocellata 

Diesing 

1850:513 

1861 

Taenia  ocellata 

Van  Beneden. 

1861 :165 

1889 

Taenia  ocellata 

Lonnberg 

1889 :14 

1902 

Ichthyotaenia  filicollis 

Schneider 

1902 :21-22 

1902 

Ichthyotaenia  ocellata 

Schneider 

1902 :23 

1903 

Ichthyotaenia  percae 

Schneider 

1903 :13-22 

1905 

Ichthyotaenia  ocellata 

Schneider 

1905 :11-15 

1905 

Ichthyotaenia  percae 

Schneider 

1905:15-17 

1911 

Proteocephalus  percae 

La  Rue 

1911 :475 

*  Specific  Diagnosis:  Characters  of  the  genus.  "Worms  of  varying 
length,  20-200  mm.  Maximum  breadth  1.1-1.5-2,0  mm.  Head  short, 
broad,  flattened  dorsoventraUy,  apex  slightly  elevated  or  flattened.  At 
summit  a  fifth  sucker,  muscular,  0.033-0.040-0.060  mm.  in  diameter. 
Four  suckers,  muscular,  with  deep  cavities,  0.085-0.100-0.137  mm.  in 
diameter,  situated  at  broadest  zone  of  head  or  immediately  anterior 
thereto.  Head  0.192-0.357  mm.  broad,  usually  about  0.300  mm.  Thick- 
ness of  head  0.170-0.238  mm.  Surface  of  head  without  furrows.  Neck 
0.170-0.50  mm.  broad  by  3.0-10.0  mm.  long.  Transition  to  first  proglot- 
tlds  imperceptible.  First  proglottids  usually  broader  than  long,  0.255- 
0.34  mm.  broad  by  0.085-0.102  mm.  long,  rarely  longer  than  broad. 
Mature  and  ripe  proglottids  broader  than  long.  Breadth  of  mature 
proglottids  0.935-1.19-1.30  mm.,  length  of  same  0.255-0.340  mm.  Breadth 
of  ripe  proglottids  1.10-1.7  mm.;  length  of  same  0.42-0.85  mm.  End 
proglottids  about  quadrate.  Proglottids  few,  150  or  more,  closely  at- 
tached. Segmentation  fairly  evident,  angles  of  proglottid  rounded. 
Surface  of  worm  wrinkled  and  rough. 

Genital  organs  as  in  genus.    Genital  aperture  marginal,  near  mid- 
dle of  proglottid,  irregularly  alternating.    Testes  in  single  layer,  irreg- 


94 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


[94 


ularly  arranged  between  vitellaria.  Testes  ovoidal,  0.05-0.095  mm.  long 
by  0.05-0.07  mm.  broad,  50-60  in  number.  Vas  deferens  a  thick  straight 
mass  of  coil$  reaching  to  middle  of  proglottid  or  beyond.  Cirrus-pouch 
0.34-0.47  mm.  long,  slender,  lying  at  right  angles  to  margin  of  proglot- 
tid. Ratio  of  length  of  cirrus-pouch  to  proglottid  breadth  1:3-2:5. 
Cirrus,  when  protruded  0.1-0.2  mm.  long,  when  unprotruded,  straight 
within  cirrus-pouch. 

Vagina  always  anterior  and  dorsal  to  cirrus-pouch,  crossing  cirrus- 
pouch  near  middle.  Vaginal  sphincter  small.  Lumen  of  vagina  ciliated. 
Ovary  bilobed,  posterior.  Lobes  long  and  heavy.  Vitellaria  lateral, 
follicular,  denser  near  ovary.  Uterus  with  4-5-9  lateral  outpocketings 
on  either  side.  Uterine  pore  single,  situated  near  middle  of  proglottid. 
Eggs  provided  with  three  membranes.  Embryo  0.01  mm.,  second  mem- 
brane 0.0264-0.029  mm.,  outer  membrane  0.031-0.037  mm.  in  diameter. 

Habitat :    Intestine  of  Perca  fluviatilis  and  other  fish. 


Host 

Locality 

Collector 

Authority 

Perca  norvegica 

Miiller 

0.  F.  MiiUer  (1788) 

Perca  fluviatilis 

Pallas 

Rudolphi  (1810) 

( 1)Acerina  cernua^ 

Pallas 

Rudolphi  (1810) 

Perca  fluviatilis 

Greifswald 

Rudolphi 

Rudolphi   (1810) 

Perca  fluviatilis 

Finland 

Schneider 

Schneider 

(1903  and  190^) 

Cor  eg  onus  lavaretus 

E]inland 

Schneider 

La  Rue 

(The  present  paper) 

Coitus  qiiadricornis^ 

Finland 

Schneider 

La  Rue 

(The  present  paper) 

Perca  fluviatilis 

Rositten,  East 
Prussia 

Miihling 

Muhling  (1898) 

{1)Gasterosteus  aculeatus^ 

Rositten,  East 
Prussia 

Miihling 

Miihling  (1898) 

Perca  fluviatilis 

Upsala 

Lonnberg 

Lonnberg  (1889) 

^In  the  course  of  this  work  specimens  from  Acerina  cernua  have  been  deter- 
mined to  belong  to  a  new  species,  Proteocephalus  cernuae.  Outwardly  these  speci- 
mens much  resemble  P.  percae  and  there  is  therefore  some  doubt  whether  the 
specimens  collected  by  Pallas  and  the  Taenia  percae  MuUer  were  identical.  Such 
a  doubt  is  valid  until  modern  investigations  show  that  Acerina  cernua  harbors  P. 
percae  (Miiller). 

^Needs  confirmation. 

^This  seems  to  be  a  misdetermination.    The  parasite  was  probably  P.  filicollis. 


9$]  PROTEOCEPHALIDAE  —  LA  RUE  95 

This  species  was  first  described  by  Miiller  (1780:152-155,  179).  His 
diagnosis  (1780:179)  reads  thus:  ''Taenia  percae,  capite  bulboso,  ocel- 
lis  quatuor,  osculis  raarginalibus,  articulis  quadrangulis. ' '  On  account  of 
its  inaccessibility  it  is  deemed  best  to  quote  here  a  portion  of  Miiller 's 
observations  (1780:152-153)  tho  in  a  measure  these  observations  are 
recapitulated  in  his  (1788)  description  which  is  quoted  after  this  of 
1780. 

"Der  Kopf  oder  das  erste  Glied  des  vordern  und  schmalen  Endes  ist  von 
gleicher  Breite  mit  den  nachsten  Gelenken,  vornc  stumpf,  und  in  der  Mitte  des 
Randes  gleichsam  eingebogen.  Oben  mit  zwei  durchsichtigen  und  iiber  einander 
stehenden  kugelrunden  Knoten  besetzt.  Wenn  der  Kopf  fast  ausgetrochnet  ist, 
zeiget  sich  die  Spur  von  vier  Zirkeln,  zwei  an  der  Stelle  der  Knoten  mit  einem 
eingedruckten  dunkeln  Punkte  in  der  Mitte;  und  zwei  gleichsam  im  Schatten  an 
der  untem  Lefze  naher  am  Rande. 

"Die  Gelenke  sind  platter,  und  kommen  einem  gleichseitigen  Quadrat  Naher, 
als  bei  den  andern  Bandwiirmern.  Auf  beiden  Seiten  gegen  den  Rand  laufen 
zwei  helle  weisse  Linien  den  ganzen  Wurm  hindurch ;  eigentlich  werden  sie,  wenn 
man  genau  zusiehet,  von  den  Verbindungen  der  Gelenke  unterbrachen,  und  machen 
in  jedem  Gelenke  eine  kleine  bogenformige  Linie;  hie  und  da  siehet  man  neben 
den  Verbindungen  der  Gelenke  in  der  Mitte  den  gewohnlichen  punktformigen 
Eindruck,  und  auf  der  einen  Flache,  nicht  auf  der  andern,  einen  weissen  Quer- 
strich,  der  sich  von  der  Mitte  des  einen  Seitenrands  bis  zu  der  Mitte  des  Gelenkes 
erstrecket.  Die  Gelenke  sind  am  Rande  dicker,  als  gewohnlich ;  viele  haben  an 
dem  einen  aussern  Seitenrande  ein  tiefes  Loch,  und,  es  scheinet  dass  der  Quefstrich 
von  dem  hier  durchfallenden  Lichte  herriihret,  weil  er  an  dem  undurchlocherten 
Gelenke,  und  an  der  Seite,  wo  kein  Loch  ist,  nicht  zu  bemerken  war.  Diese 
Locher  sind  wahre  Vertiefungen,  und  immer  in  der  Mitte  des  Seitenrandes  der 
Gelenke,  doch  nicht  in  alien,  sondern  ohne  Ordnung,  bald  in  zwei,  bald  in  mehreren 
auf  einander  folgenden.  Bei  den  andern  Bandwiirmern  habe  ich  dergleichen  nicht 
bemerket." 

Under  the  name  Taenia  cystica  Pallas  (1781:101)  described  some 
plerocercoids  which  he  had  found  encysted  in  the  liver  of  the  perch  and 
pike.  It  seems  probable  that  the  specimens  from  the  liver  of  the  perch 
were  the  larval  form  of  the  Taenia  percae.  It  is  improbable  that  the 
larvae  found  in  the  liver  of  the  pike  belonged  to  this  species.  Rudolphi 
(1802a  :112)  states  that  these  were  probably  Tricuspidaria.  Batsch 
(1786:156-7)  wrote  concerning  "Der  Leberbandwurm  in  Barschen". 
Under  this  title  he  considered  the  larvae  encysted  in  the  livers  of  the 
pike  and  two  species  of  perch  which  he  called  *Stock-und  Kaulbarschen'. 
His  statement  "die  Haken  und  Kopfe"  etc.  shows  very  conclusively 
that  he  was  dealing  with  some  other  species  than  Taenia  percae  tho  some 
of  his  specimens  may  have  belonged  to  that  species.     Batsch    (1786: 


96  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [96 

234-5)  compiled  a  description  of  Taenia  percae  from  Miiller's  (1780) 
description. 

Miiller  (1788:5)  gave  a  description  and  diagnosis  together  with 
drawings  of  this  species.  The  latter  have  been  faithfully  redrawn  and 
are  reproduced  (Figs.  121,  122,  172,  173).  These  are  of  value  for  com- 
parative purposes  while  the  description  which  is  here  quoted  is  suf- 
ficiently detailed  to  permit  its  comparison  with  later  descriptions.  His 
diagnosis  and  synonymy  are  omitted. 

"Caput,  seu  extremitas  antica  et  tenuior  insequentium  articulorum  latitudine, 
obtusum  supra  bulbis  binis  sphaericis,  ocularibus,  pellucentibus,  subtus  binis  simili- 
bus  obsoletis  infra  marginem  instructum.  Articuli  corporis  crassiores  et  planiores 
quam  in  congeneribus  figuram  quadrangulam  imitantur.  In  ipso  corpora  linea 
longitudinalis  Candida  utrinque  conspicitur;  haec  propius  inspecta  e  lineolis 
articulorum  arcuatis  quavis  intersectione  interruptis  componitur.  In  ipsis  intersec- 
tionibus  porus  seu  osculum  solitum  passim  conspicitur,  ac  marginis  medio  in  altera 
paginae  parts,  non  in  opposita,  linea  transversa  alba  ad  medium  paginae  pertingens 
pellucet.  Haec  foramini  in  ipsissima  ora  laterali  quorundam  articulorum  conspicuo 
ac  ad  medium  usque  pertuso  -deberi  videtur.  Foramen  hoc  canaliculatum  oviductus 
est  ac  in  paucis  Taeniae  speciebus  observatur. 

"In  paene  exsiccato  capitulo  quatuor  circuli  bulborum  vestigia  apparuere; 
posteriores  puncto  opaco  impressi  erant. 

"In  intestinis  Percae  marinae  raro;  ultra  \4ginti  enim  diversa  aetate  examini 
subieci,  in  duabus  tantum  Martio  et  Aprili,  unicam  nempe  in  altera,  in  altera 
tres  reperi. 

"Quas  Claris.  GOEZE  in  Siluro  reperit,  sola  absentia  lineolarum  lateralium 
non  in  omnibus  aeque  visibilium  diflferre  videtur." 

Miiller  in  his  synonymy  gave  Taenia  alternatim  transverse  lineata 
Goeze,  a  parasite  of  Silurus  glanis,  as  a  synonym  of  Taenia  percae. 
This  was  followed  by  Gmelin  (1790:3079)  who  also  considered  Goeze 's 
species  to  be  the  same  as  Taenia  percae.  Schrank  (1788:48)  in  his 
catalogue  gave  a  diagnosis  of  Taenia  percae  and  listed  the  perch  as  the 
host.  Rudolphi  (1802a-112-13)  diagnosed  and  described  this  species, 
naming  it  Taenia  ocellata.  He  recognized  that  his  species  and  Miiller's 
Taenia  percae,  exclusive  of  the  variety  /?,  were  synonymous.  His 
description  is  here  quoted  verbatim : 

"Taenia  ocellata :  capite  obtuso,  osculis  orbicularibus  excavatis ;  corpore  plan*- 
iusculo,  articulis  subquadrangulis. 

"Taenia  percae  Syst.  Nat.  p.  3079.  n.  77.  (exclusa  var.  3.) 
"Zwei  bis  ftinf  Zoll  lang.  Der  Kopf  rundlich,  der  Gestalt  nach  alle  Augen- 
blicke  verschieden,  bald  aufgeblasen,  bald  zusammengezogen  u.  s.  w.  Nach  vorne 
stehen  am  Kopf  vier  kleine  aber  tiefe  Saugblasen,  wie  Napfschen,  an  denen  ich 
keinen  Rand,  wie  bei  der  vorigen  Art,  wahrgenommen  habe ;  sonst  sind  sie  eben 
so  v«randerlich.  Der  Hals  ist  dunn  und  schwach  runzlich,  die  auf  ihn  folgenden 
Glieder  werden  allmahlich  gfrosser  und  grosser,  so  dass  die  Starksten  derselben  fast 


97]  PROTEOCEPHALIDAE  —  LA  RUE  97 

eine  Linie  breit  sind.  In  der  Mitte  eines  jeden  grossern  Gliedes  erscheinen  Linien 
die  sich  untereinander  verbinden  und  so  fast  das  ganze  died  zuletzt  einnehmen, 
doch  habe  ich  keine  Miindungen  an  den  Randern  wahrnehmen  konnen.  Die 
Glieder  sind  deutlich  und  beinahe  viereckig,  man  kann  also  nicht  leicht  diesen 
Wurm  mit  der  Triciispidaria  verwechseln,  wenn  man  auch  vom  Kopfende  absehen 
wollte,  das  sonst  schon  allein  statt  alles  Unterschiedes  ist. 

"Ich  habe  diesen  Wurm,  aber  nur  selten,  im  Darmkanal  des  Barsches,  Perca 
fluviatilis,  gefunden,  und  zwar  im  Junius.  Pallas,  der  diesen  Wurm  unstreitig 
gekannt  hat,  (N.  Nord,  Beitr.  I.  S.  102.  (113)  Fig.  33.  A.)  verwechselt  ihn  mit 
der  Tricuspidaria,  da  er  ihn  nicht  allein  im  Darmkanal  sondern  auch  in  Blasen  an 
der  Leber  beim  Barsch  (und  Hecht)  gefunden  haben  will;  in  diesen  Blasen 
namlich  ist  er  wohl  nicht  zu  finden. 

"Miillers  Taenia  percae  (Zool.  Dan.  II.  p.  5.  Tab.  44.  Fig.  1-4.)  scheint 
hierher  zu  gehoren,  der  Kopf  (bis  auf  dessen  Spitze,  dergleichen  ich  nicht  finde), 
passt  gut;  was  er  vom  Korper  sagt,  bezeichnet  sehr  grosse  Exemplare,  dergleichen 
auch  Fig.  I.  abbildet,  ich  kann  also  aus  den  Oeffnungen  u.  s.  w.  keinen  Grund 
degegen  hernehmen.  Eins  aber  verstehe  ich  nicht,  er  sagt  namlich :  Articuli  corporis 
crassiores  et  planiores  quam  in  congeneribus ;  ich  finde  sie  auch  etwas  dick 
und  abgeplattet,  aber  wenn  sie  dick  sind,  konnen  sie  naturlich  nicht  so  fiach  seyn,  als 
bey  den  ganz  diinnen  Arten.  Uebrigens  ist  der  Bandwurm,  den  Muller  beschreibt, 
aus  der  Perca  marina. 

"Wenn  aber  Muller  und  auf  seine  Auctoritat  auch  Gmelin  den  Bandwurm 
aus  dem  Wels  dahin  ziehen  will,  so  kann  ich  nicht  ihrer  Meinung  seyn.  Sie 
kennen  jenen  Wurm  nur  aus  Goezes  Abbildung,  und  die  ist,  besonders  wenn  man 
den  Kopf  betrachtet,   durchaus  verschieden." 

Zeder  (1803:355)  gave  a  diagnosis  and  synonymy  of  this  species 
under  the  name  Holy  sis  percae.  He  remarked :  * '  Hier  sind  in  Gmelins 
Ausgabe  versehiedene  Arten  zusammengeworfen.  Naeh  dem  Zitat  des 
Pallas  (N.  Bord.  Beytr.  S.  102)  hat  Gmelin  den  Blasen  wurm  aus  der 
Leber  des  Bersehen  hieher,  und  zwar  gans  irrig,  gerechnet.  Und  dass  der 
aus  dem  Darmkanal  des  Welses  nicht  hieher  gehoret,  erhellet  schon  aus 
den  Bestimmungen  desselben."  Rudolphi  (1810:108)  again  diagnosed 
and  described  Taenia  ocellata.  His  synonymy  is  almost  identical  with  the 
one  which  has  been  given  up  to  this  point  in  this  work  with  the  exception 
that  he  considered  Taenia  cernuae  to  be  a  synonym  of  Taenia  ocellata. 
His  habitat  data  are  here  quoted:  "In  intestinis  Percae  fluviatilis  et 
cernuae  Pallas  copiose,  Percae  marinae  Muller  rarius,  repererunt.  Ipse 
vario  anni  tempore  in  Perca  fluv.  copiosam  offendi. ' '  He  considered  that 
the  variety  ^  in  Gmelin 's  (1790:3079)  catalogue  (Taenia  alternatim 
transverse  lineata  Goeze)  belonged  with  Taenia  osculata  and  not  here. 
Rudolphi  (1819:149)  quoted  his  earlier  diagnosis  (1810).  He  further 
stated  that  Taenia  ocellata  had  been  found  abundantly  by  himself  in 
Perca  fluviatilis  at  Greifswald,  by  Pallas  in  Perca  cernua,  and  by  Miil- 
ler  rarely  in  Perca  norvegica,  a  fish  which  Miiller  called  Perca  marina. 


98  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [98 

Perca  cernua  is  now  known  as  Acerina  cernua  while  the  Perca 
marina  should  be  known  as  Sehastes  marina,  according  to  Jordan  and 
Evermann  (1896-1900).  Rudolphi  (1819)  stated  that  Perca  marina  did 
not  occur  in  Miiller's  region  and  that  the  latter 's  specimens  came  from 
Perca  norvegica.  It  seems  very  doubtful  that  the  species  designated 
by  Miiller  as  Perca  marina  and  by  Rudolphi  as  Perca  norvegica  can  be 
identical  with  Sehastes  marina,  for  the  latter  is  a  marine  fish  whereas 
the  host  of  Taenia  percae  must  be  a  freshwater  fish  since  the  genus 
Proteocephalus  occurs  only  in  the  fishes  of  freshwater.  Dujardin 
(1845:583)  and  Diesing  (1850:513)  added  almost  nothing  to  the  data 
as  given  by  the  earlier  investigators. 

Van  Beneden  (1861:165)  briefly  reported  Taenia  ocellata  but  gave 
no  drawings.  He  stated  that  the  perch  was  the  host.  Von  Linstow 
(1878:208,  209)  did  not  increase  the  list  of  hosts.  Zschokke  (1884: 
16-17)  reported  and  described  a  cestode  from  Perca  fluviatilis.  Lake 
Lucerne  which  he  identified  as  Taenia  filicolUs  Bud.  As  shown  in  an- 
other part  of  this  work  his  specimens  are  probably  to  be  considered  as 
belonging  to  the  species,  P.  duhius.  Under  the  name  of  Taenia  ocel- 
lata Zschokke  (1884:13,  14)  described  a  form  from  several  hosts  includ- 
ing Coregonus  fera.  It  seems  not  at  all  unlikely  that  this  species  is  the 
same  as  the  one  from  Coregonus  fera  which  Kraemer  (1892)  first  de- 
scribed as  Taenia  filicolUs  and  later  in  the  same  article  as  Taenia 
ocellata.  Kraemer 's  species  has  been  shown  elsewhere  in  this  mono- 
graph to  be  very  different  from  Taenia  percae,  and  it  was  described 
by  La  Bue  (1911)  as  a  new  species,  P.  fallax.  A  more  complete  discus- 
sion of  Zschokke 's  and  Kraemer 's  work  above  cited  will  be  found  in  the 
description  and  synonymy  of?,  fallax.  Lonnberg  (1889:14)  reported 
Taenia  ocellata  Bud.  from  Perca  fluviatillis,  Upsala.  His  diagnosis  is 
short,  yet  sufficient  data  are  given  to  enable  one  to  determine  that  this 
form  is  not  the  same  as  that  one  which  he  reported  (1889:15)  from 
Gasterosteus  pungitius  and  identified  by  him  as  Taenia  filicollis  Bud. 
Lonnberg 's  Taenia  ocellata  was  nearly  3  mm.  broad  by  150  mm.  long. 
It  is  highly  probable  that  this  form  is  identical  with  the  Taenia  percae 
Miiller.  Linton  (1897:425-426)  provisionally  assigned  to  this  species 
some  cestodes  which  in  all  probability  belong  to  Proteocephalus  amhlo- 
plitis  (Leidy).  Von  Batz  (1897:453)  reported  finding  Ichthyotaenia 
ocellata  in  Esox  lucius  and  Lucioperca  sandra  in  Lake  Balaton.  It  seems 
improbable  that  his  determination  was  correct.  Miihling  (1898:36) 
found  what  he  determined  to  be  Ichthyotaenia  ocellata  in  Perca  fluvia- 
iUis  and  Gasterosteus  aculeatus  at  Bositten,  East  Prussia.  It  seems 
probable  that  he  has  made  a  misdetermination  in  the  case  of  the  cestodes 
found  in  Gasterosteus  aculeatus. 


99]  PROTEOCEPHAUDAE  —  LA  RUE  99 

Schneider  (19p2:21-22)  reported  a  species  of  cestode  parasitic  in 
Perca  fluviatilis  which  he  considered  to  be  Ichthyotaenia  filicollis.  He 
believed  this  to  belong  to  the  species  found  by  Zschokke  (1884)  in  the 
same  host  species.  He  noted  the  fact  that  Kraemer  (1892)  considering 
his  Taenia  filicollis  to  be  a  younger  stage  of  Taenia  ocellata,  put  his 
two  forms  together  in  the  same  species,  T.  ocellata.  To  this  last 
Schneider  did  not  agree  but  held  that  the  specimens  from  Perca  fluvi- 
atilis and  Coregonus  lavaretus  were  distinct  species.  He  noted  on  the 
one  hand  the  near  relationships  and  the  great  similarities  of  the  species 
of  the  Ichthyotaenia  and  on  the  other  extreme  variability  of  the  cestodes. 
He  concluded  that  it  was  not  therefore  wholly  unlikely  that  the  one 
form  stands  in  relation  to  the  other  as  a  variety  which  has  arisen  from 
the  changed  environmental  conditions  of  the  new  host.  He  further  gave 
a  short  description  of  the  worm  in  which  he  considered  external  charac- 
ters almost  exclusively.  In  the  same  article  Schneider  (1902:23)  re- 
ported some  specimens  of  Ichthyotaenia  ocellata  from  Coregonus 
lavaretus  and  from  Cottus  quadricornis.  It  seems  improbable  that  the 
latter  species  is  a  host  of  the  cestode  mentioned. 

Schneider  (1903:13-23)  decided  to  avoid  entirely  the  questions  of 
identity  surrounding  the  names  Taenia  ocellata  and  Taenia  filicollis  Rud. 
He  proposed  to  use  the  older  name  Taenia  percae  to  designate  that  form 
from  Perca  fluviatilis  which  he  (1902:21-22)  had  previously  considered 
to  be  Taenia  filicollis  Rud.  His  reasons  for  this  action  need  not  be 
stated  here.  As  synonyms  of  Ichthyotaenia  percae  he  cited  Taenia 
percae  MiiUer  (1788:5,  pi.  XLIV),  Taenia  ocellata  Rudolphi  (1810:108), 
Taenia  filicollis  Zschokke  (1884:16-18)  in  part,  Ichthyotaenia  ocellata 
Riggenbach  (1896:268)  in  part,  and  Ichthyotaenia  filicollis  Schneider 
(1902:21)  in  part.  It  is  to  be  noted  that  Schneider  failed  to  include 
in  his  synonymy  any  part  of  Kraemer 's  (1892)  Taenia  ocellata.  This 
seems  the  more  remarkable  when  it  is  remembered  that  both  Kraemer 
and  Riggenbach  were  students  under  Zschokke  and  it  is  quite  likely 
that  both  the  pupils  accepted  the  determination  of  their  master  on  the 
latter 's  T.  filicollis.  Nevertheless  parts  of  Zschokke 's  and  Riggenbach 's 
data  on  T.  filicollis  are  held  to  apply  to  Ichthyotaenia  percae.  Nor  did 
Schneider  refer  to  Benedict  (1900)  in  his  synonymy.  In  this  he  was 
correct  for  in  another  part  of  this  work  it  is  shown  that  Benedict 
was  working  on  a  distinct  species  for  which  La  Rue  (1911)  has  pro- 
posed the  name  Proteocephalus  exiguus.  In  his  later  description  of  the 
species  Schneider  (1905)  gave  nothing  further  on  the  synonymy  of 
Ichthyotaenia  percae  but  he  added  some  descriptive  data  to  that  given 
in  his  papers  of  1902  and  1903. 

Liihe  (1909)  considered  Taenia  ocellata  Rud.  a  sjnQonym  of  Ichthyo- 
taenia percae  (0.  F.  MiiUer).    In  his  diagnosis  he  followed  Kraemer 


100  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [100 

(1892)  and  as  an  illustration  of  the  proglottid  of  the  species  he  appar- 
ently redrew  Kraemer's  figure  18.  Apparently  also  he  either  ignored 
Schneider's  work  or  else  he  considered  Kraemer's  T.  ocellata  and 
Schneider's  Ichthyotaenia  percae  to  be  identical  without  having  made  a 
comparison  of  the  material.  No  one  who  has  made  a  careful  comparison 
of  Schneider's  and  Kraemer's  drawings  and  descriptions  would  think 
of  putting  them  in  the  same  species.  It  has  been  shown  elsewhere  in 
this  monograph  that  Kraemer's  T.  ocellata,  or  T.  filicoUis  as  he  called 
his  species  in  one  part  of  his  paper,  is  not  the  same  as  Schneider's 
Ichthyotaenia  percae.  Kraemer's  specimens  doubtless  belong  to  Proteo- 
cephalus  fallax  La  Bue. 

A  comparison  of  Miiller's  (1780  and  1788)  data  and  drawings  of 
T<i€nia  percae  with  Rudolphi's  (1802a  and  1810)  diagnoses  and  descrip- 
tions of  Taenia  ocellata  compels  the  writer  to  believe  that  they  were 
referring  to  the  same  form.  It  is  also  to  be  remembered  that  Rudolphi 
gave  Miiller's  Taenia  percae  as  a  synonym  of  his  own  T.  ocellata.  There 
remains  no  reason  why  the  name  which  Miiller  suggested  for  this  species 
should  not  be  used  instead  of  the  name  suggested  by  Rudolphi  many 
years  later.  The  generic  name  is  Proteocephalus.  Attention  is  called 
to  the  fact  that  Miiller's  figure  1  (reproduced  Fig.  173)  compares  very 
favorably  with  Schneider's  figures  and  with  his  specimens  which  the 
writer  has  examined.  Miiller's  figure  4  (reproduced  Pig.  172)  shows 
the  shape  of  the  proglottids  and  the  location  of  the  genital  organs  very 
well.  The  light  area  extending  from  the  margin  nearly  to  the  middle 
of  the  segment  is  doubtless  the  cirrus-pouch  which  under  certain  cir- 
cumstances could  have  been  observed  by  Miiller.  Miiller's  figures  2  and 
3  of  the  heads  (reproduced*  Fig.  121,  122)  are  much  like  the  heads  of 
Schneider's  specimens  which  the  writer  has  examined.  Taken  all  in 
aU  the  identification  seems  to  be  as  complete  as  is  possible  without  a 
study  of  Miiller's  and  Rudolphi's  actual  specimens. 

Since  the  time  of  Rudolphi  the  specific  names  Taenia  filicoUis  and 
T.  ocellata  have  appeared  frequently  in  the  literature  of  helminthology. 
For  the  most  part  the  names  are  listed  in  reports  of  parasites  found, 
without  diagnosis  and  without  description.  In  such  cases  the  identity 
of  the  forms  so  named  can  not  be  determined  with  any  degree  of 
accuracy.  A  consideration  of  the  host  can  at  times  throw  a  little  light 
on  the  subject  but  this  datum  alone  is  not  to  be  trusted.  It  is  noteworthy 
that  in  the  case  of  Taenia  filicoUis  no  author  prior  to  Zschokke  claimed 
to  find  this  species  in  any  other  than  the  sticklebacks,  and,  indeed,  in 
only  two  species  of  sticklebacks.  Zschokke  (1884)  reported  this  species 
from  a  number  of  hosts,  none  of  them  sticklebacks.  In  the  case  of 
Taenia  ocellata  no  author  prior  to  Zschokke  reported  this  species  in  any 


101]  PROTEOCEPHALIDAE—LA  RUE  101 

other  species  than  Perca  fluviatUis,  P.  norvegica,  and  Acerina  cernua. 
Zschokke  (1884)  reported  this  species  from  several  hosts,  and  from  the 
same  host  species  he  reported  several  other  species  of  Taenia.  A  number 
of  years  later  Kraemer,  a  pupil  of  Zschokke,  attempted  to  prove  that 
Taenia  filicollis  was  the  young  stage  of  Taenia  ocellata,  and  since  the 
appearance  of  Kraemer 's  work  (1892)  Taenia  filicollis  and  Taenia 
ocellata  have  for  the  most  part  been  considered  to  be  identical.  As  a 
result  of  the  tacit  acceptance  of  this  determination  great  confusion 
exists  in  the  identification  of  many  lots  of  specimens.  It  is  therefore 
difficult  or,  indeed,  quite  impossible  to  tell  what  references  among  the 
later  writers  should  be  considered  in  the  synonymy  of  Proteocephalus 
percae.  An  actual  study  of  many  of  the  specimens  bearing  these  much 
debated  names  must  be  made  before  an  exact  determination  is  possible. 
A  comparative  study  of  specimens  of  this  genus  now  to  be  found  in  the 
helminthological  collections  in  Europe  would  doubtless  yield  many  inter- 
esting results.  In  the  preparation  of  this  monograph  five  lots  of  material 
identified  as  Taenia  ocellata  Rud.  have  been  studied.  Four  of  these 
lots  were  received  from  European  investigators,  another  from  an 
American.  Out  of  these  five  lots  four  have  proved  to  be  new  species 
reported  by  La  Rue  (1911)  while  the  fifth  was  Proteocephalus  percae. 
La  Rue  (1911:475)  gave  this  species  a  place  in  a  list  of  Proteocephalus 
species. 

Professor  "Ward  secured  from  Professor  Levander,  Helsingfors, 
Finland,  some  specimens  of  Schneider's  species,  Ichthyotaenia  percae 
(0.  F,  Miiller)  and  /.  ocellata  (Rud.)  This  material  included  both 
alcoholics  and  slides.  One  bottle,  now  No.  10.123  in  Professor  Ward's  col- 
lection, bears  the  original  label:  "Ichthyotaenia  percae  0.  F.  M.  Bet. 
G.  Schneider."  A  second  bottle  of  this  material,  now  No.  10.122,  in  Pro- 
fessor Ward's  collection,  bears  the  original  label:  '* Ichthyotaenia  ocel- 
lata Rud.  Coregonus  lavaretus.  14,  V,  02. ' '  The  two  slides  were  labelled 
*' Ichthyotaenia  ocellata,  Coregonus,  24,  VIII,  01."  From  this  data  it 
seems  that  these  must  have  been  prepared  from  the  same  lot  which 
Schneider  mentioned  in  his  report  (1902:23  and  53).  Sections  and  toto 
preparations  were  made  from  specimens  of  lot  No.  10.123,  while  sections 
only  were  prepared  from  lot  No.  10.122.  Some  of  the  heads  of  each  lot 
were  cleared  in  glycerine. 

A  comparison  of  Schneider's  descriptions  of  his  Ichthyotaenia 
percae  and  his  7.  ocellata  show  but  two  significant  differences  between 
the  species.  He  states  that  the  head  of  I.  percae  is  much  the  larger  but 
that  the  main  difference  between  them  is  in  the  presence  in  the  head 
of  7.  ocellata  of  "einen  fiinften  flachen  Saugnapf  von  40/li  im  Durch- 
raesser  an  der  Spitze,  der  offenbar  auch  noch  funktioniert,  da  er  aus 


102  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [102 

einer  grossen  Anzahl  von  Radialmuskelfasern  besteht".  Of  this  organ 
in  /.  percae  he  writes,  * '  an  der  Spitze  des  Scolex  findet  sich  das  deutliehe 
Scheitel-organ,  welches  bei  konservierten  Exemplaren  eine  kugelformige 
Anhaufung  in  der  Langsaxe  des  Tieres  gestrekter  Zellen  darstellt,  die 
ihren  Charakter  als  Muskel-zellen  offenbar  vorloren  haben.  Wir  sehen 
hier  ein  rudimentares  Organ,  das,  wie  es  scheint,  jede  Funktion  einge- 
biisst  hat,"  The  other  differences  in  size  and  proportion  of  heads  and 
necks  may  be  explained  as  being  due  to  contraction  states.  The  breadth 
of  the  head  when  but  a  very  few  specimens  can  be  measured  is  a  valuable 
tho  not  an  absolutely  reliable  diagnostic  character  in  a  form  which  is  so 
variable  and  so  contractile.  In  measuring  Schneider's  own  specimens 
the  writer  found  the  heads  of  the  two  forms  very  much  alike.  One  head 
of  the  /.  percae  was  even  smaller  than  the  dimension  recorded  by 
Schneider  for  I.  ocellata,  while  every  head  of  I.  ocellata  was  broader  than 
the  dimensions  which  he  secured  from  measurements  of  the  same  form. 
Their  thickness  in  the  writer's  measurements  was  about  equal  to  the 
breadth  as  he  recorded  it.  It  is  evident  that  size  of  head  is  not  a  valid 
character  for  the  differentiation  of  these  two  forms. 

As  for  the  fifth  sucker,  examination  of  frontal  sections  of  heads  of 
I.  percae  with  an  oil  immersion  lens  revealed  the  true  muscular  structure 
of  a  sucker.  The  drawing  (Fig.  73)  shows  the  nuclei,  the  radial  muscles, 
basement  membrane,  cut  ends  of  circular  muscle  fibers,  and  the  cuticula 
covering  the  surface  of  the  sucker.  No  cavity  was  noted  in  the  fifth 
sucker  of  either  form.  This  last  difference  then  between  the  two  forms 
vanishes.  An  examination  of  the  two  lots  of  Schneider's  specimens 
revealed  a  marked  agreement  in  every  diagnostic  feature  except  in  the 
proportions  of  the  proglottid  s  and  this  slight  difference  was  undoubtedly 
due  to  different  states  of  contraction.  Most  remarkable  similarity  was 
found  in  the  relations  of  the  cirrus,  cirrus-pouch,  vas  deferens,  testes, 
vagina,  vaginal  sphincter,  uterus,  vitellaria,  eggs  and  position  of  the 
genital  pore.  An  examination  of  the  comparative  table  and  of  the 
drawings  (Figs.  8,  73,  74,  69,  70)  of  preparations  from  Kraemer's  7. 
ocellata  and  drawings  (Figs.  9,  71,  72)  of  his  /.  percae  will  at  once  show 
the  strong  similarities.  The  differences  which  Schneider  noted  in  the 
shape  of  the  ovary  are  readily  explained  as  being  due  to  contraction 
states  of  the  proglottid.  Schneider's  I.  ocellata  is  clearly  identical  with 
his  /.  percae,  and  on  account  of  the  priority  of  the  name  suggested  by 
Miiller  the  latter  should  be  retained.  It  should  also  be  clearly  recognized 
that  Taenia  ocellata  is  a  synonym  of  Taenia  percae  and  hence  can  not  be 
used  to  designate  any  other  species  of  cestodes  in  this  genus.  Schneider 
then  was  not  justified  in  using  the  name  Taenia  ocellata  for  the  cestode 
found  in  Coregonus  lavaretus  even  had  it  proved  to  be  a  new  species. 


103]  PROTEOCEPHALIDAE—LA  RUE  103 

These  cestodes  vary  in  length  from  20  to  200  mm.  and  their  maxi- 
mum breadth  varies  from  1.1  to  1.5  to  2.0  mm.  Differences  in  the  stage 
of  development  and  also  in  the  amount  of  contraction  account  for  the 
greater  part  of  this  variation.  The  short  broad  head  (Figs.  8,  9,  121, 
122)  is  flattened  dorsoventrally.  Its  apex  may  be  slightly  elevated  or  it 
may  be  somewhat  flattened.  At  the  summit  is  a  small  but  muscular  fifth 
sucker.  The  four  suckers,  directed  forward  and  outward,  are  situated 
at  the  broadest  zone  of  the  head  or  just  anterior  thereto.  The  breadth 
of  the  head  varies  from  0.192  to  0.357  mm.  tho  the  greater  number  of 
heads  measured  0.30-0.34  mm.  In  thickness  the  head  measures  0.170- 
0.238  mm.  Such  variations  in  dimensions  are  due  to  the  states  of  con- 
traction of  the  muscles  of  the  neck  and  head.  Strong  contractions  of  the 
longitudinal  muscles  of  the  neck  and  head  cause  the  neck  to  dilate 
markedly.  This  condition  also  causes  the  posterior  parts  of  the  head  to 
widen  and  thicken,  and  thus  the  suckers  are  directed  more  nearly  for- 
ward. The  converse  of  these  statements  also  holds.  The  relaxed  condi- 
tion of  the  longitudinal  muscles  and  the  contracted  condition  of  the  neck 
muscles  result  in  the  narrow  head.  The  surface  of  the  head  is  smooth, 
without  wrinkles  or  furrows.  The  four  muscular  suckers  measure  0.085- 
0.100-0.137  mm.,  the  fifth  sucker  0.033-0.06  mm.  in  diameter.  The 
surface  of  the  neck  is  smooth  tho  at  times  transverse  wrinkles  simulate 
proglottids.  The  neck,  usually  thin,  varies  considerably  in  length  and 
breadth.  It  measures  0.170-0.050  mm.  broad  while  its  length  varies  from 
3.0  to  10.0  mm.  Its  transition  to  the  first  proglottids  is  scarcely  per- 
ceptible. 

First  proglottids  are  extremely  variable  in  shape.  In  a  strongly 
contracted  worm  they  are  much  broader  than  long,  being  about  0.34  mm. 
broad  by  0.085  mm.  long.  In  less  contracted  individuals  these  first  pro- 
glottids may  measure  0.255  mm.  broad  by  0.102  mm.  long.  Schneider 
(1903)  states  that  they  are  broader  than  long,  nearly  quadrate  or  longer 
than  broad.  Mature  and  ripe  proglottids  show  somewhat  similar  var- 
iations tho  in  less  degree.  Mature  proglottids  in  all  specimens  examined 
by  me  were  broader  than  long.  The  breadth  varied  from  0.935  to  1.19 
or  even  as  much  as  1.30  mm.,  and  the  length  from  0.255  to  0.340  mm. 
Fully  ripe  proglottids  are  slightly  longer  in  proportion  to  breadth.  They 
measure  1.1  mm.  broad  by  0.42  mm.  long  and  in  the  case  of  the  largest 
proglottids  1.7  mm.  broad  by  0.85  mm.  long.  No  end-proglottid  was  ob- 
served by  me.  Schneider  describes  such  a  segment  as  being  about  quad- 
rate. The  number  of  segments  varies  from  the  few  in  the  very  short 
worms  to  150  or  more  in  the  longer  specimens  (Schneider).  Segmenta- 
tion is  fairly  evident  especially  when  the  worm  is  viewed  with  a  lens  of 
low  magnification,  for  the  angles  of  the  proglottids  are  rounded  and  the 


104  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [104 

intersegmental  furrows  are  plainly  marked.  Shallow  grooves  and  de- 
pressions give  the  surface  of  the  worm  a  somewhat  roughened  appear- 
ance.   The  segments  are  firmly  attached  to  each  other. 

A  common  genital  aperture  (Figs.  69,  70,  71,  72,  172)  is  situated 
near,  or  slightly  anterior  to,  the  middle  of  the  margin  of  each  proglottid. 
Its  position  in  the  strobila  alternates  irregularly  from  side  to  side.  There 
is  no  genital  papilla. 

The  testes  (Figs.  69,  72)  lie  in  a  single  layer  between  the  vitellaria 
and  anterior  to  the  ovary.  They  are  of  ovoidal  or  spheroidal  shape.  In 
length  and  breadth  they  measure  0.053-0.080-0.095  mm.  by  0.05-0.07  mm. 
In  Schneider's  drawing  of  his  I.  ocellata  50-60  testes  may  be  seen,  and 
in  preparations  from  either  lot  of  his  material  about  that  number  of 
testes  may  be  counted.  He  stated  that  the  testes  measure  0.10  by  0.06 
mm.  The  vas  deferens  forms  a  thick  straight  mass  of  coils  extending 
from  the  end  of  the  cirrus-pouch  to  the  middle  of  the  segment  or  even 
a  little  beyond.  In  mature  and  ripe  proglottids  the  vas  deferens  is 
always  well  distended  with  spermatozoa.  Extending  into  the  segment  at 
right  angles  to  the  margin  is  the  long  and  slender  cirrus-pouch.  This, 
in  a  reconstruction  (Fig.  74)  from  a  transection,  may  be  seen  to  curve 
upward  to  the  dorsal  wall  of  the  dermo-muscular  sac  where  it  is  firmly 
attached  by  strong  muscle  fibers.  Its  length  varies  somewhat  in  pro- 
glottids of  different  stages  of  development  and  in  different  stages  of 
contraction.  It  measures  0.34-0.37-0.425-0.47  mm.  long,  the  greater 
lengths  occurring  in  greatly  contracted,  and  hence  very  wide,  proglottids. 
Schneider  reported  the  ratio  of  its  length  to  the  proglottid  breadth  as 
1 :3  in  his  /.  ocellata  and  1 :3rl  :2  in  his  I.  percae.  The  writer  finds  the 
ratio  in  each  of  his  forms  to  be  from  1 :3  to  2 :5.  In  no  case  did  the  cirrus- 
pouch  reach  to  the  middle  of  the  proglottid  tho  at  first  sight  the  coils  of 
the  vas  deferens  frequently  gave  it  that  appearance.  Schneider  saw  the 
protruded  cirrus  in  7.  percae,  where  it  measured  0.1-0.2  mm.  In  both 
of  his  forms  he  found  the  cirrus  straight  in  the  cirrus-pouch.  The  same 
condition  was  observed  by  the  writer.  Schneider  did  not  record  having 
seen  a  j)rotruded  cirrus  and  the  writer  failed  to  find  such  cirri  in  the 
material  examined.  From  the  fact  that  the  cirrus  and  the  ductus 
ejaculatorius  form  a  straight  tube  in  the  cirrus-pouch  one  can  safely 
postulate  that  the  protruded  cirrus  would  be  short. 

The  vagina  always  opens  dorsal  to  the  cirrus-pouch.  This  is  shown 
by  a  reconstructed  transection  (Fig.  74)  through  this  region.  Then  pass- 
ing inward  and  bending  slightly  anteriad  it  crosses  the  cirrus-pouch  ob- 
liquely near  the  middle  and  passes  toward  the  ventral  wall  of  the  dermo- 
muscular  sac.    From  the  point  of  crossing  the  cirrus-pouch  (Fig.  72) 


105]  PROTEOCEPHALIDAE—LA  RUE  105 

it  passes  posteriad  in  a  long  curve  to  the  interovarial  space.  The  vagina 
lies  just  dorsal  to  the  uterus.  Very  near  the  opening  of  the  vagina  a 
short  sphincter  vaginae  0.020-0.026  mm.  thick  may  be  seen.  The  inner 
surface  of  the  vagina  is  apparently  ciliated.  A  receptaculura  seminis  has 
not  been  found,  tho  it  may  exist.  The  ovary  (Figs.  69,  70,  72)  is  long 
and  heavy.  It  is  somewhat  arched  in  the  more  elongated  proglottids  and 
more  elongated  and  slender  in  the  longitudinally  contracted  proglottids. 
Frontal  sections  show  the  ovary  to  be  made  up  of  closely  connected 
tubes  or  branches.  The  vitellaria  are  long  lateral  follicular  glands  which 
extend  from  the  anterior  end  of  the  segment  to  the  ovary  but  not  beyond. 
The  vitellaria  are  more  dense  near  the  ovary. 

The  coils  of  the  vagina,  uterine  passage,  oviduct,  and  the  common 
vitelline  duct  nearly  fill  the  interovarial  space.  Here  also  are  the  oocapt 
and  ootype.  The  relations  and  connections  of  these  various  passages  are 
similar  to  those  described  by  Benedict  (1900)  for  this  group.  An  ill- 
defined  uterine  passage  discharges  into  the  uterus  near  the  middle  of  the 
segment.  The  uterus  (Figs.  70,  71)  in  ripe  proglottids  has  4,  5,  7,  8  or 
even  9  lateral  outpocketings  on  either  side.  These  come  to  fill  up  nearly 
the  whole  ventral  side  of  the  segment.  Schneider  found  but  a  single 
uterine  pore  near  the  middle  of  the  proglottid.  The  writer  has  found 
this  number  to  be  correct  for  his  specimens.  Schneider  measured 
the  eggs  as  follows:  embryo  0.025  mm.;  second  membrane  0.045-0.050 
mm.;  outer  membrane  0.090-0.125  mm.  My  measurements  are  made 
from  uterine  eggs  of  alcoholic  materials.  The  writer  has  already  shown 
(La  Rue  1909)  that  the  outer  membrane  swells  up  when  it  comes  in  con- 
tact with  the  water.  It  is  not  certain  that  the  middle  one  does  this.  The 
writer's  measurements  of  eggs  from  Schneider's  material  are  as  follows: 
embryo  alone  0.019;  second  membrane,  0.0264-0.029;  outer  membrane, 
0.031-0.037  mm.  Since  the  writer  has  had  Schneider's  own  specimens 
for  examination  he  is  inclined  to  believe  that  Schneider  measured  the 
embryo  plus  its  investing  membrane.  The  eggs  which  the  writer  so 
measured  gave  his  figures  almost  exactly  for  the  diameter  of  the  embryo. 
It  is  a  matter  of  regret  that  there  is  not  more  strict  unanimity  in  the 
method  of  measuring  and  recording  the  measurements  of  cestode  eggs. 
The  data  comprised  in  the  discussion  of  the  preceeding  pages  regarding 
P.  oscellatus  and  P.  percae  are  presented  in  tabular  form  on  the  next 
page. 


106 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


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108  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [108 

An  examination  of  the  preeeeding  table  makes  evident  that  the 
differences  between  Schneider's  Proteocephalus  percae  and  his  P.  oceU 
latus  are  so  slight  that  one  would  not  be  justified  in  considering  these 
forms  as  varieties.  Schneider  dealt  with  a  single  species  which  ac- 
cording to  the  rule  of  priority  should  be  known  as  Proteocephalus  percae 
(MiiUer). 

Proteocephalus  percae  in  some  respects  resembles  P.  agonis  Bar- 
bieri,  P.  exiguus  La  Rue  and  P.  pusillus  Ward.  It,  however,  is  so  much 
larger  than  these  species  that  the  possibility  of  an  identity  is  entirely 
precluded.  P.  percae  in  many  ways  resembles  P.  fallax  La  Rue  but  it 
is  larger  than  that  species.  The  character  of  its  proglottids  is  greatly 
different.  P.  percae  has  many  more  and  larger  testes,  a  longer  cirrus- 
pouch,  a  much  larger  ovary,  more  voluminous  vitellaria,  a  larger  head, 
larger  suckers,  and  larger  sucker  openings.  P.  fallax  has  a  larger  num- 
ber of  preformed  uterine  pores.  It  also  has  larger  embryos.  The  area 
occupied  by  the  ovaries  of  ripe  proglottids  is  triangular  in  P.  fallax  but 
much  elongated  in  P.  percae.  P.  percae  resembles  P.  cernuae  in  the 
size  of  the  head,  in  the  size  and  character  of  the  fijth  sucker,  in  the  shape 
of  the  ovary,  and  somewhat  in  the  character  of  the  proglottids.  P. 
percae,  however,  has  larger  suckers  with  larger  sucker  openings,  a  longer 
and  more  slender  neck,  a  much  longer  cirrus-pouch,  and  fewer  and 
smaller  testes.  The  relationship  of  cirrus-pouch  and  vagina  are  greatly 
different  in  the  two  species.  In  P.  percae  the  vagina  crosses  the  cirrus- 
pouch  near  the  middle  of  the  latter  while  in  P.  cernuae  the  vagina  does 
not  cross  the  cirrus-pouch.  P.  cernuae  may  have  more  uterine  pouches 
and  its  embryos  are  larger  than  in  P.  percae.  Further  points  of  dif- 
ference between  P.  percae  and  P.  cernuae  may  be  readily  noted  in  an 
examination  of  the  comparative  table  of  Proteocephalus  species  (vide 
infra). 

P.  percae  in  some  respects  resembles  P.  longicolUs  as  described  by 
von  Linstow  but  it  differs  in  the  number  of  uterine  pouches,  number  of 
testes,  length  of  cirrus-pouch,  a'nd  in  the  size  of  eggs  and  of  suckers. 
P.  percae  most  closely  resembles  P.  pinguis  La  Rue.  The  heads  of  the 
two  species  are  of  about  the  same  size.  The  suckers  are  much  alike  but 
the  fifth  sucker  is  much  better  developed  in  P.  pinguis  than  in  P.  percae. 
P.  pinguis  is  more  slender,  its  proglottids  more  nearly  quadrate  and 
thicker  than  in  P.  percae.  P.  pinguis  has  many  more  uterine  pouches 
and  preformed  uterine  pores.  Its  cirrus-pouch  is  less  than  half  as  long 
as  that  of  P.  percae.  P.  percae  is  readily  differentiated  from  P.  torulosus 
by  reason  of  the  total  lack  of  a  fifth  sucker  in  the  latter.  In  number  of 
uterine  pouches,  length  of  cirrus-pouch  and  in  numerous  other  ways  these 
species  are  very  dissimilar. 


109]  PROTEOCEPHALIDAE—LA  RUE  109 


PROTEOCEPHALUS  LONGICOLLIS    (Zeder) 
[Figs.  167-169] 


1780: 

Taenia  eperlani 

Acharius 

1780 :3080 

1789: 

Taenia  salmonis  Wartmanni 

Frolich 

1789 :24 

1790: 

Taenia  Froelichii 

Gmelin 

1790 :3080 

1800: 

Alyselminthus  longicollis 

Zeder 

1800:258. 

1802: 

Taenia  longicollis 

Rudolphi 

1802a  :113 

1803: 

Halysis  longicollis 

Zeder 

1803 :333 

1803: 

Taenia  Renkina 

Schranli 

1803 :242 

1810: 

Taenia  longicollis 

Rudolplii 

1810:107 

1810: 

Scolex  tetrastomus 

Rudolplii 

1810:6 

1819: 

Taenia  longicollis 

Rudolplii 

1819 :149 

1845: 

Taenia  longicollis 

Dujardin 

1845 :585 

1850: 

Taenia  longicollis 

Diesing 

1850 :512-513 

1891: 

Taenia  longicollis 

V.  Linstow 

1891 :565-576 

1891: 

Tetracotylus  longicollis 

IVIonticelli 

1891 :162 

1894: 

Ichthyotaenia  longicollis 

Lonnberg 

1894:803 

1911: 

Proteocephalus  longicollis 

La  Rue 

1911 :475 

Specific  Diagnosis:  Characters  of  genus.  Cestodes  of  small  size. 
Observed  length  as  much  as  20-200-464  mm.,  breadth  1-2-2.25  mm. 
Scolex  0.43  mm.  broad,  length (?).  Necls:  0.3  mm.  broad  by  3.2  mm. 
long.  Suckers  circular,  0.12-0.14-0.18-0.19  mm.  in  diameter.  Fifth 
sucker  half  as  large  as  others.  First  proglottids  0.43  mm.  long  by  0.71 
mm.  broad.  Proglottids  at  end  of  first  third  of  strobila  0.53  mm.  long 
by  0.99  mm.  broad.  Posterior  proglottids  1.03  mm.  long  by  0.83  mm. 
broad.     End  proglottid  triangular. 

Genital  pore  marginal,  irregularly  alternating,  situated  at  end  of 
anterior  one-fourth  of  proglottid.  Testes  spheroidal,  0.09  mm.  in  diam- 
eter, 25  in  each  segment,  arranged  in  two(  ?)  fields  near  vitellaria.  Vas 
deferens  forming  a  close  mass  in  the  median  field  of  proglottid.  Cir- 
rus-pouch spindle-shaped,  muscular,  0.110  mm.  broad,  extending  just 
through  vitellaria.  Cirrus  club-shaped  when  everted,  reaching  0.2  mm. 
past  margin  of  segment.  Ovary  bilobed,  posterior.  Lobes  slender, 
united  by  a  midpiece.  Vitellaria  lateral,  follicular.  Uterus  with  three 
lateral  pouches  on  either  side.  Vagina  anterior  to  cirrus-pouch,  not 
crossing  same.  Sphincter  vaginae?  Receptaculum  seminis?  Unripe 
eggs  0.0156-0.0196  mm.  in  diameter. 


110 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


[110 


Habitat:    In  intestine  of  host. 


Host 


Coregonus  (Saliiio)  lavaretus 
Qoregonus  {Saltno)  maraenula 

=  C.  albula 
Coregonus  (Salmo)  wartmann't 
Coregonus  (Salmo)  wartmanni 
Coregonus  wartmanni  nobilis 
Coregonus  fera 
Coregonus  fera 

Coregonus  albula 

Coregonus  schinzii  helveticus 

Coregonus  exiguus  albellus 

Osmerus  eperlanus 

Osmerus  eperlanus 

Osmerus  eperlanus 

Trutta  trutta 

Trutta  trutta 

Trutta  fario 

Salmo   thymallus^  Thymallus 

vulgaris 
Salmo   thymallus  =  Thymallus 

vulgaris 
Salmo  alpinus 
Salmo  salvelinus 
Salmo  salvelinus 
Salmo  umbla 
Esox  lucius 
Esox  lucius 
Perca  fluviatilis 
Alburnus  lucidus 
Squalius  cephalus 
Squalius  leuciscus 


Locality 


Greifswald 
Berlin 


Lake  Lucerne 
Lake  Geneva 
Rositten,  East 

Prussia 
Nicolaiken, 

East  Prussia 
Lake  Lucerne 
Lake  Lucerne 


Berlin 


(in  Museum 
Vienna) 


Lake  Lucerne 
Lake  Geneva 
Lake  Balaton 
Lake  Lucerne 
Lake  Lucerne 
Lake  Lucerne 
Lake  Lucerne 
Lake  Lucerne 


Collector 


Rudolphi 
Rudolphi 

Frolich 

Schrank 

Nufer 

Zschokke 

Miihling 

Miihling 

Nufer 

Nufer 

Acharius 

Rudolphi 

V.  Linstow 

Frolich 

Zeder 

Martin 

Rudolphi 

Bremser 

Martin 


Nufer 

Zschokke 

von  Ratz 

Nufer 

Nufer 

Nufer 

Nufer 

Nufer 


Authority 


Rudolphi, 
Rudolphi, 

Rudolphi, 

Rudolphi, 

Nufer, 

Zschokke, 

Miihling, 


1810:107. 
1810:107. 

1810:107. 
1819:495. 
1905 :75 
1884:11,14-16. 
1898 :36. 


Miihling,      1898 :36. 


Nufer, 

Nufer, 

Rudolphi, 

Rudolphi, 

v.  Linstow, 

Rudolphi, 

Rudolphi, 

Rudolphi, 

Rudolphi, 


1905  75. 
1905  75. 
1810 :2i2. 
1810:107, 
1891 :56s-576. 
1810:107. 
1810:107. 
1810:107. 
1810:107. 


Rudolphi,     1819 :49s. 


Rudolphi, 

v.  Linstow, 

Nufer, 

Zschokke, 

von   Ratz, 

Nufer, 

Nufer, 

Nufer, 

Nufer, 

Nufer, 


1810:107. 
1878 :262. 
1905  75- 
1884:11,14-16. 
1897:159. 
1905  75- 
190S  75. 
1905  75. 
1905  75. 
1905  75- 


Possibly  this  species  is  the  form  which  was  originally  referred  to 
by  Acharius  (1780:52)  who  called  it  Taenia  eperlani.  Acharius 's  de- 
scription and  diagnosis  of  this  form  are  not  accessible  to  the  writer. 
Despite  the  fact  that  Rudolphi  (1810:212)  considered  that  this  species 
could  not  have  been  a  synonym  of  Taenia  longicollis  Rud.  it  seems  that  it 
must  be  a  Synonym  of  the  latter  species  or  else  some  of  the  host  records  of 


Ill]  PROTEOCEPHALIDAE—LA  RUE  111 

Rudolphi  and  others  are  open  to  question.    Concerning  this  species  Ru- 
dolphi  (1810:212)  wrote: 

"In  Salmonis  Eperlani  cavo  abdominis  Taeniam  reperit  (pro  Fasciola  habi- 
tam)  quatuor  ad  quinque  lineas  longam,  capite  oblongo  obtuso;  osculis  duobus 
superioribus  anticis  (totidem  inferioribus  in  figura  latentibus)  orbicularibus  exi- 
guis ;  collo  nuUo ;  articulis  transversis,  obtusis.  Cum  collum  nullum  delineatum 
sistatur,  pro  Taenia  longicolli  (n.  20)  eodem  in  pisce  obvia  vix  haberi  potest,  huic 
enim  longissimum  conceditur." 

Frolich  (1789:24)  named  a  species  Taenia  salmonis  Wartmanni 
which  is  recognized  by  Rudolphi  ( 1802a  :113  and  1810 :107 )  as  a  synonym 
of  Taenia  longicollis  Rud.  Frolich 's  description,  if  he  gave  one,  is  not 
accessible  to  the  writer.  The  name  with  which  he  designated  the  species 
is  unavailable  on  account  of  its  trinomial  character.  Gmelin  (1790: 
3080)  gave  a  diagnosis  of  Taenia  Froelichii.  This  species  was  consid- 
ered by  Rudolphi  (1802a  :113  and  1810:107)  to  be  a  synonym  of  Taenia 
longicollis.  Gmelin 's  diagnosis  reads:  ''TAENIA  FROELICHII.  No. 
91. — T.  capite  cum  collo  longissimo  tenuissimo  marticulato  continuo,  ova- 
riis  dendaticis  linea  laterali  cinctis.  Froelich  Naturf.  24  p.  124.  t.  4.  /. 
20.  21.    Habitat  in  Salmonis  Wartmanni  intestino  duodeno." 

The  name  longicollis  as  a  designation  for  a  species  of  cestode  was 
first  used  by  Zeder  (1800:258)  in  connection  with  the  generic  name 
Alyselminthus.  Unfortunately  Zeder 's  description  is  not  available  to 
the  writer  who  therefore  can  not  judge  as  to  its  character.  Rudolphi 
(1802a  :113)  gave  a  specific  diagnosis  of  this  species  which  he  styled 
Taenia  longicollis.  In  his  synonymy  he  cited  Alyselminthus  longicollis 
Zeder,  Taenia  Salmonis  Wartmanni  Froelich,  and  Taenia  Froelichii 
Gmelin.    After  a  statement  regarding  the  hosts  of  this  form  he  wrote : 

"Zeders  Beschreibung  ist  voUkommen  geniigend;  wenn  er  Frolich  tadelt,  weil 
dieser  die  baumschenartigen  Figuren  oder  Eyerstocke  der  Glieder  mit  zur  Bestim- 
mung  des  Wurms  braucht,  so  habe  ich  im  AUgemeinen  nichts  dagegen,  dass  man 
sie  weglasst.  Bey  einigen  Arten  sind  sie  indessen  besonders  ausgezeichnet,  so 
auch  hier,  wo  man  sie  auf  den  ersten  Blick  gewahr  wird,  wesgegen  auch  die 
letzten  Glieder  blaulich,  oft  sogar  schwarzlich  aussehen.  Wo  man  erst  den  Press- 
schieber  zu  Hiilfe  nehmen  muss,  um  sie  zu  sehen,  darf  man  ihrer  nicht  erwahnen, 
das  versteht  sich." 

Rudolphi  gave  no  description  of  this  species  at  this  time  nor  were 
there  any  drawings  to  accompany  his  comments.  Zeder  (1803:333)  re- 
ferred to  this  species  under  the  name  Halysis  longicollis.  In  his  syn- 
onymy he  lists  Taenia  salmonis  Wartmanni  Frolich,  Alyselminthus  lon- 
gicollis Zeder,  Taenia  Frolichii  Gmelin.    His  diagnosis  reads :    * '  HALY- 


y^ 


112  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [112 

SIS  LONGICOLLIS. — leviter  crenata;  capite  cum  collo  longissimo,  lin- 
eari  continuo,  tnmcato;  osculorum  margine  prominente." 

Kudolphi  (1810:107)  gave  a  diagnosis  and  description  which  is 
here  quoted  verbatim: 

"Taenia  longicollis  R. 

Taenia:  capite  truncato,  collo  longissimo,  articulis  subquadratis,  ovariis  race- 
mo  sis. 

Frolich  im  Naturforsch.  24.P.124.  Tab4.  fig.20.21.  Taenia  Salmonis  Wart- 
manni. 

Gmel.  Syst.  Nat.  p.  3080.  n.  91.    Taenia  Froelichii. 

Zeder  Nachtrag  p.  258.     Alyselminthus  longicollis. 

Rudolphi  in  Wied.  Arch.  III.  i.p.113.    Taenia  longic. 

Zeder  Naturg.  p.  333.  n.  9.     Halysis  longic. 

"Hab.  in  intestinis  Salmonum.  In  Salmonibus  Wartmanni  plurimis  Froelich- 
ius  Aug^sto  copiosam;  in  5".  Trutta  Zederus;  ego  in  5".  Lavareto  Majo  et  S.  Ma- 
raenulis  plurimis  Februario,  reperimus.  In  Salmone  Eperlano  a  se  inventas  anx 
Treviranus  mecum  communicavit. 

"Descr.  Vermes  unum  ad  septum  pollices  longi,  lineam  dimidiam  vel  integram 
lati. 

"Caput  depressum,  truncatum,  exiguum;  osculis  orbicularibus,  binis  tam  su- 
perioribus,  quam  inferioribus,  capitis  margini  antico  approximatis,  ut  sub  ejusdem 
motu  saepe  antica  omniaque  simul  appareant.  Colluni  cum  capite  continuum,  lon- 
gissimum,  tertiam  circiter  totius  longitudinis  partem  sibi  vindicans,  depressum, 
margine  obtuso  integerrimo.  Corpus  depressum,  articulus  anterioribus  brevissimis, 
reliquis  subquadratis,  marginibus  singulorum  antico  et  postico  rectis,  lateralibus 
rotundatis,  ut  totius  vermis  latera  crenata  appareant.  Articulus  ultimus  obtusus. 
Ovaria  in  articulis  posticis,  inde  vel  caerulescentibus,  vel  nigrescentibus,  distincta, 
racemosa  sive  dendritica. 

"Obs.  I.  An  scolex  tetrastomus  supra  dictus  hujus  Taenia  proles  nondum 
articulata?  Sed  reliquarum  Taeniarum  foetus  semper  articulates  vidi,  neque  caput, 
neque  pars  postica  acuta  conveniunt. 

"Obs.  2.  Hujus  speciei  cum  insequentibus  ob  articulos  margine  postico  vix 
incumbentes,  ob  capitis  collique  formam,  affinitas  magna;  differentiae  tamen  spe- 
cificae  singularum  discrimen  satis  indicant. 

"Obs.  3.  An  quas  Stellerus  (Pallas  N.  Nord.  Beytr.  I.i, p,i02)  in  Eperlani 
vesica  natatoria  reperit,  Taenia  hue  pertinent?     Conf.  n.  113." 

Kudolphi  (1819:149)  quoted  the  diagnosis  used  by  him  before  (Ru- 
dolphi, 1810:107).  He  stated  that  he  had  found  the  species  in  Corego- 
nus  (Salmo)  lavaretus  at  Greifswald,  in  C.  (Salmo)  maraenula=C.  aU 
hula  and  in  Osmerus  eperlanus  at  Berlin,  that  Frolich  had  found  it  in 
Coregonus  (Salmo)  wartmanni,  Frolich  and  Zeder  in  Trutta  trutta. 
Specimens  in  the  museum  at  Vienna  had  been  found  in  Salmo  thymal- 
lus=Thymallus  vulgaris.  He  further  stated  that  Martin  had  found  it 
in  Salmo  alpinus  and  in  Trutta  fario.    This  serves  as  a  very  excellent 


113]  PROTEOCEPHALIDAE—LA  RUE  113 

summary  of  the  hosts  which  had  been  recorded  for  this  species  up  to 

this  time.    Rudolphi  (1819:495)  stated  that  Bremser's  specimens  which 

had  been  taken  in  the  pyloric  caeca  of  Salmo  thymallus  belonged  to 

Taenia  longicollis  and  not  to  Bothriocephalus.    He  further  stated  that 

Taenia  Renkina  Schrank  is  identical  with  Taenia    longicollis    because  1"^ 

Salmo  Wartmanni  is  called  Renken   in    central    Germany.     Schrank 's 

(1803 :242)   description  of  Taenia  renkina  is  not  at  hand  hence  it  is 

necessary  to  follow  Rudolphi's    judgment    in    the    matter.     Dujardin 

(1845:585)  added  but  little  to  the  data  of  the  earlier  authors.    Diesing 

(1850:512-13)  gave  a  literature  review  that  is  of  value.    Von  Linstow 

(1878:262)  in  his  catalogue  of  entozoa  gave  Salmo  salvelinus  as  a  host 

of  this  species. 

Zschokke  (1884:11,  14-16)  identified  and  described  some  cestodes 
from  Coregonus  fera  and  Salmo  umhla,  Lake  Geneva,  as  Taenia  longi- 
collis. A  careful  comparison  of  his  descriptions  of  T.  longicollis  and  T. 
ocellata,  likewise  reported  by  him  from  the  same  hosts,  causes  the  writer 
to  conclude  that  his  specimens  belonged  to  the  same  species.  Length 
forms  the  chief  difference  between  his  species.  In  describing  certain 
organs  of  the  two  forms  he  used  almost  identical  phrases.  Kraemer 
(1892)  in  his  study  of  Taenia  ocellata  evidently  used  length  as  the  chief 
distinguishing  character.  Zschokke 's  description  of  T.  longicollis  does 
not  agree  with  the  description  and  figures  of  that  form  as  given  by  von 
Linstow  (1891:565-576).  Von  Linstow 's  specimens  were  taken  from 
Coregonus  eperlanus,  one  of  the  hosts  in  which  Rudolphi  found  T.  lon- 
gicollis. It  is  therefore  probable  that  von  Linstow 's  specimens  rather 
than  Zschokke 's  belong  to  Rudolphi 's  species.  Moreover,  von  Linstow 's 
description  agrees  more  completely  with  Rudolphi 's  than  does  Zschok- 
ke's.  Lonnberg  (1894:803)  included  Taenia  longicollis  Rud.  in  the  list 
of  species  in  his  genus  Ichthyotaenia. 

Zschokke  (1896:772-777)  listed  Ichthyotaenia  longicollis  only  in 
Trutta  fario  from  Lake  Geneva.  Thanks  to  Prof.  H.  B.  "Ward,  the 
writer  has  been  able  to  examine  parts  of  a  strobila  of  Zschokke 's  **T. 
longicollis  aus  Forelle"  (Trutta  fario)  and  is  able  to  state  positively 
that  it  is  not  the  same  species  as  von  Linstow 's  T.  longicollis.  Nor  is  it 
the  same  as  the  specimens  from  Coregonus  fera  which  Zschokke  sent  to 
Professor  Ward  as  T.  ocellata  (No.  0.99  in  Prof.  H.  B.  Ward's  collec- 
tion, the  type  of  P.  fallax  La  Rue).  Von  Ratz  (1897:159)  listed  Ich- 
thyotaenia longicollis  from  Esox  lucius  from  Lake  Balaton  in  Hungary. 
This  is  probably  a  misdetermination.  Von  Ratz  gave  no  description  so 
not  even  a  probable  determination  can  be  made.  Miihling  (1898:36) 
reported  Ichthyotaenia  longicollis  Rud.  from  Coregonus  alhula  at  Nico- 
laiken  and  from  Osmerus  eperlanus  at  Rositten  in  East  Prussia.     He 


114  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [114 

gave  no  description.  Nufer  (1905:75)  in  the  report  of  an  investigation 
of  Lake  Lucerne  reported  Proteocephalus  longicollis  from  Perca  fluvia- 
tilis,  Alhurnus  lucidus,  Squalius  cephalus,  S.  leuciscus,  Esox  lucius, 
Coregonus  wartmanni  nohilis,  C.  exiguus  albellus,  C.  schinzii  helveticus 
and  Salmo  salvelinus.  It  is  extremely  doubtful  if  the  specimens  of  Pro- 
teocephalus which  Nufer  found  in  other  than  the  salmonoid  fishes  be- 
long to  the  species  P.  longicollis.  Nufer 's  tabulated  description  of  P. 
longicollis  (Nufer  1905:147)  is  based  almost  entirely  upon  the  work  of 
von  Linstow  (1891).  The  character  of  Nufer 's  work  has  been  discussed 
at  some  length  in  connection  with  P.  macrocephalus  and  P.  torulosus. 
Many  of  the  statements  made  in  those  places  regarding  his  work  apply 
here. 

A  discussion  of  the  facts  brought  out  in  this  historical  summary 
cannot  yield  very  satisfactory  conclusions.  For  the  most  part  the  au- 
thors cited  have  necessarily  been  compelled  to  depend  upon  the  exter- 
nal features  of  the  worm  for  diagnostic  characters.  Benedict  (1900) 
showed  how  little  dependence  could  be  placed  on  such  characters  alone 
for  descriptive  purposes.  The  earlier  workers  necessarily  based  their 
conclusions  on  little  else  than  external  features,  upon  records  of  hosts, 
and  locality  of  collection.  The  host  records  of  all  the  workers  who 
reported  this  species  prior  to  Nufer  and  von  Ratz  show  that  these  men 
regarded  this  species  as  being  peculiar  to  the  salmonoid  fishes  and  it 
seems  quite  probable  that  they  were  correct  in  this  respect.  It  is  im- 
possible to  determine  whether  this  species  is  parasitic  in  all  the  salmon- 
oid fishes  of  the  list  or  whether  a  number  of  cestode  species  have  been 
reported  under  the  one  name  without  a  painstaking  comparative  study 
of  such  specimens. as  exist  in  private  and  museum  collections  together 
with  a  study  of  specimens  from  hosts  and  localities  as  indicated  in  the 
list  of  hosts.  It  is  not  likely  that  this  will  be  done,  at  least  not  for  some 
time. 

The  question  of  priority  of  name  is  not  an  important  one.  This 
species  is  known  as  one  of  Eudolphi's  species  probably  on  account  of 
the  weight  of  Rudolphi's  authority  as  much  as  on  anything  else.  His 
first  notes  on  this  species  contain  only  the  briefest  diagnosis  and  he  dis- 
tinctly says,  "Zeders  Beschreibung  ist  vollkommen  geniigend",  etc,  etc 
(vide  supra).  In  the  light  of  this  it  seems  that  Zeder  should  be  cred- 
ited with  the  specific  name  longicollis.  It  is  true  of  course  that  Rudolphi 
was  the  first  to  use  the  combination  Taenia  longicollis.  As  to  the  use 
of  the  name  longicollis  or  some  of  the  names  that  were  proposed  still 
earlier,  viz..  Taenia  Froelichii,  T.  salmonis  wartmanni  and  T.  eperlani, 
it  may  be  said  that  the  identity  of  the  last  is  not  well  known.  Ru- 
dolphi (1819)  thought  it  was  not  identical  with  T.  longicollis.    T.  ml- 


115]  PROTEOCEPHALIDAE—LA  RUE  115 

monis  Wartmanni  being  a  descriptive  name  of  three  parts  is  not  avail- 
able. It  is  not  necessary  to  discuss  the  question  as  to  whether  it  was 
sufficiently  described  to  give  it  any  standing.  T.  Froelichii  was  evi- 
dently based  on  the  same  description  as  the  T.  salmonis  Wartmanni  for 
Gmelin  (1790)  gives  little  more  than  a  catalogue  of  species  and  refers 
to  Froelich's  (1789)  work.  Since  the  combination  Taenia  longicollis 
Rudolphi  is  no  longer  the  proper  one  with  which  to  designate  this 
species,  the  generic  name  Proteocephalus  being  used  to  designate  the 
genus,  it  is  here  suggested  that  the  proper  combination  for  the  designa- 
tion of  the  species  is  Proteocephalus  longicollis  (Zeder).  In  a  recent 
article  the  writer,  La  Rue  (1911:475),  overlooked  the  priority  of  Ze- 
der's  Alyselminthus  longicollis  over  Rudolphi 's  Taenia  longicollis  and 
hence  in  a  list  of  species  of  Proteocephalus  gave  the  credit  for  the  P. 
longicollis  to  Rudolphi  instead  of  Zeder. 

The  description  is  based  on  von  Linstow's  (1891)  paper.  His  ma- 
terial was  collected  from  Osmerus  eperlanus  which  is  one  of  the  hosts 
in  which  Rudolphi  found  Taenia  longicollis. 

Von  Linstow  reports  that  his  largest  specimen  measured  46.4  mm. 
long  by  a  maximum  breadth  of  0.99  mm.  At  its  posterior  end  the  worm 
measured  0.83  mm.  broad.  The  scolex  measured  0.43  mm.  broad,  the 
neck  0.3  mm.  broad  by  3.2  mm.  long,  first  proglottids  0.43  mm.  long  by 
0.71  mm.  broad,  proglottids  at  end  of  first  third  of  strobila  0.53  mm. 
long  by  0.99  mm.  broad.  The  proglottids  gradually  increase  in  length 
until  the  last  are  1.03  mm.  long  by  0.83  mm.  broad.  The  end-proglottid 
is  triangular  and  drawn  out  posteriorly.  The  specimen  was  incom- 
pletely developed.  The  four  suckers  have  a  circular  outline,  0.12-0.14- 
0.18-0.19  mm.  in  diameter.  A  fifth  apical  sucker  has  a  diameter  equal 
to  one-half  that  of  the  others. 

The  excretory  system  is  made  up  of  two  large  lateral  vessels,  one 
on  either  side  just  outside  of  the  vitellaria,  and  six  smaller  vessels,  three 
on  either  side  lying  dorsal  to  the  larger  vessel.  All  excretory  vessels 
are  greatly  twisted  and  many  anastomoses  connect  them.  About  0.08 
mm.  anterior  to  the  posterior  margin  of  each  proglottid  a  transverse 
commissure  connects  the  ventral  excretory  vessels.  The  latter  measure 
0.019  in  diameter  and  the  smaller  vessels  about  0.0078  mm.  In  the 
region  just  posterior  to  the  suckers  there  is  a  circular  commissure  of 
excretory  vessels  from  which  branches  extend  anteriorly  into  the  head. 
All  the  main  vessels  discharge  into  a  small  vesicle  at  the  posterior  end 
of  the  end-proglottid. 

The  genital  pore  is  marginal,  irregularly  alternating,  and  situated 
near  the  end  of  the  anterior  one-fourth  of  the  proglottid  (Fig.  167). 
The  testes  are  large,  spheroidal,  measuring  up  to  0.09  mm.  in  diameter. 


116  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [116 

Von  Linstow  states  that  there  are  about  25  testes  in  each  segment, 
and  that  these  are  situated  in  two  fields  near  the  vitellaria.  In  a  draw- 
ing of  a  transverse  section,  reproduced  (Fig.  168),  von  Linstow  figures 
the  testes  in  two  layers.  This  causes  the  writer  to  suspect  that  the 
number  of  testes  (25)  as  stated  by  von  Linstow  is  too  small.  In  all  the 
species  of  Proteocephalus  studied  by  the  writer  the  testes  have  been  in 
one  layer  if  the  number  of  testes  was  small  and  never  in  two  layers 
unless  they  numbered  at  least  fifty.  Frequently  one  sees  proglottids 
containing  more  than  fifty  testes  lying  in  a  single  layer.  It  should  also 
be  borne  in  mind  that  von  Linstow  figured  ten  testes  in  his  cross-section 
of  the  proglottid.  Ten  out  of  twenty-five  is  too  large  a  proportion  of 
testes  to  appear  in  one  thin  section  through  proglottids  of  this  size.  One 
wonders  what  was  left  for  the  other  sections  of  which  there  must  have 
been  quite  a  number.  Since  the  proglottid  which  von  Linstow  delin- 
eates is  ripe  or  at  least  contained  eggs  in  the  uterus  it  seems  probable 
that  he  overlooked  the  testes  of  the  median  region  where  the  eggs  would 
be  densely  packed.  In  the  transverse  section  to  which  reference  has 
been  made  the  region  dorsal  to  the  coils  of  vas  deferens  is  free  from 
testes,  a  condition  which  exists  in  all  the  other  species  of  the  genus 
whether  the  number  of  the  testes  be  few  or  many.  If  von  Linstow 's 
statement  that  the  testes  lie  in  two  lateral  fields  be  correct  then  this 
species  is  the  only  exception  to  the  rule  that  the  testes  in  this  genus  are 
irregularly  scattered  in  the  field  between  the  lateral  vitellaria.  The 
considerations  above  presented  cause  one  to  doubt  the  accuracy  of  von 
Linstow 's  statements  regarding  the  number  and  position  of  the  testes. 
The  vas  deferens  forms  a, thick  mass  of  coils  in  the  middle  of  the  pro- 
glottid. This  mass  apparently  does  not  extend  over  to  the  cirrus- 
pouch.  In  von  Linstow 's  figure  the  cirrus-pouch,  which  he  describes 
as  a  spindle-shaped  muscular  organ  with  a  breadth  of  0.110  mm.,  ex- 
tends just  a  little  within  the  vitellaria,  or  about  V^-Ys  across  the  pro- 
glottid. The  cirrus  (Fig.  169)  in  the  46.4  mm.  specimen  was  visible  at 
a  point  13.8  mm.  posterior  to  the  head.  It  is  short  and  club-shaped,  0.34 
mm.  long,  and  it  extends  0.2  mm.  beyond  the  proglottid 's  margin. 

A  comparison  of  cirrus-pouches  in  Von  Linstow 's  figures  (repro- 
duced Figs.  167  and  169)  shows  some  evident  discrepancies.  The  cirrus- 
pouch  in  Figure  167  is  set  far  within  the  tissues  of  the  proglottid  and  it 
is  connected  with  the  exterior  by  a  slender  tube.  Figure  169  shows  by 
far  the  more  typical  condition  and  is  to  be  considered  as  the  normal 
for  this  species. 

The  bilobed  ovary  lies  in  the  posterior  part  of  the  proglottid.  The 
lobes  are  club-shaped.  The  vitellaria  lie  in  the  lateral  fields.  They  dis- 
charge the  yolk-cells  through  the  paired  vitelline  ducts  which  pass  to 


117]  PROTEOCEPHALIDAE—LA  RUE  117 

the  interovarial  space  where  they  unite  to  form  a  common  duct  that 
empties  into  the  ootyp^.  Thus  far  von  Linstow  is  correct  in  his  descrip- 
tion of  the  organs  of  the  interovarial  space.  According  to  him  the  va- 
gina discharges  into  the  ootype  as  do  also  the  two  lobes  of  the  ovary. 
He  provides  no  visible  means  for  the  escape  of  the  fertilized  eggs  from 
the  ootype.  The  writer  agrees  with  Monticelli  (1891:162)  in  thinking 
that  von  Linstow  has  probably  mistaken  the  oocapt  for  the  ootype,  and 
has  erroneously  figured  the  vitelline  ducts  which  pass  near  the  oocapt 
and  the  shell-glands  lying  near  as  discharging  into  it.  The  barrel-like 
form  of  the  ootype  as  he  figures  it  (Fig.  167)  is  much  more  typical  of 
the  oocapt.  The  ootype  has  its  long  axis  lying  lengthwise  of  the  oviduct 
and  not  perpendicular  to  it.  The  vagina  in  von  Linstow 's  figure  is 
doubtless  the  oviduct.  There  is  every  reason  to  believe  that  the  organs 
of  the  interovarial  space  bear  the  same  relations  to  each  other  in  this 
species  as  they  do  in  the  other  members  of  the  genus.  The  vagina  opens 
into  the  common  genital  sinus  always  anterior  to  the  cirrus-pouch. 
Without  crossing  the  latter  the  vagina  describes  a  curved  course  to  the 
interovarial  space  where  it  forms  several  coils.  Von  Linstow  did  not 
mention  the  presence  of  a  sphincter  vaginae.  The  uterus  has  three 
large  lateral  pouches  on  either  side.  A  uterine  pore  is  lacking.  The 
latter  could  perhaps  be  found  after  a  careful  examination.  The  unripe 
eggs  measure  0.0156-0.0196  mm.  in  diameter.  The  plerocercoid  of  this 
species  is  found  in  the  liver  of  the  host  in  which  the  adult  is  found. 

It  is  evident  from  a  study  of  Von  Linstow 's  figures  and  text  that  he 
made  several  misinterpretations  of  the  structural  plan  of  this  cestode. 
He  recognized  that  its  plan  differed  in  certain  respects  from  the  plan 
on  which  the  Taenias  are  formed  for  he  said  that  the  Taenias  of  fish 
formed  a  distinct  group  in  the  genus  Taenia.  He  apparently  did  not 
note  the  marked  resemblance  of  their  plan  to  the  plan  of  the  Tetra- 
phyllideans.  Whether  his  failure  to  recognize  the  relationships  of  this 
species  was  the  cause  or  the  effect  of  his  misinterpretation  of  its  struc- 
ture cannot  be  determined.  At  any  rate  because  of  some  doubtful  points 
his  material  should  be  re-examined  and  a  comparative  study  made  if 
such  be  possible. 

This  species  is  readily  separated  from  the  P.  torulosus  by  its  fifth 
sucker,  and  by  its  somewhat  smaller  head  and  smaller  suckers.  The 
ovarian  lobes  are  more  slender  in  this  species  than  in  P.  torulosus.  P. 
longicollis  is  readily  differentiated  from  P.  percae  by  its  larger  head, 
larger  suckers,  shorter  cirrus-pouch,  fewer  (?)  testes,  fewer  uterine 
pouches  and  by  the  position  of  the  genital  pore.  P.  longicollis  resembles 
P.  cernuae  in  a  very  few  particulars  such  as  shortness  of  cirrus-pouch. 
However,  P.  longicollis  has  a  larger  head,  larger  suckers,  fewer  uterine 


118 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


[lis 


pouches,  smaller  embryos,  fewer  and  smaller  testes  which  are  differently 
arranged.  P.  longicoUis  is  much  larger  than  P.  fallax  La  Kue  and  P. 
dubius  La  Rue  and  it  differs  further  in  having  fewer  uterine  pouches, 
a  shorter  cirrus-pouch,  fewer  ( ?)  and  larger  testes.  The  genital  pore 
is  further  anteriad  in  P.  longicoUis  than  in  P.  fallax  or  P.  dubius.  P. 
longicoUis  differs  from  P.  dubius  La  Rue  in  having  fewer  uterine 
pouches,  a  shorter  cirrus-pouch,  fewer  and  larger  testes  and  a  different 
arrangement  of  the  same.  P.  longicoUis  does  not  closely  resemble  any 
species  reported  from  North  America  up  to  the  present  time. 


PROTEOCEPHALUS  TORULOSUS  (Batsch) 
[Figs.  7,  78-80,  145,  146,  184] 

1782 :  Taenia  articulis  rotundis 

1786:  Taenia  torulosa 

1788:  Taenia  orbicularis 

1791 :  Taenia  simplex 

1790 :  Taenia  torulosa 

1795 :  Taenia  cyprini  idi 

1800:  Bhytelminthus  cyprini 

1802:  Taenia  torulosa 

1803 :  Halysis  torulosa 

1810:  Taenia  torulosa 

1819:  Taenia  torulosa 

1845:  Taenia  torulosa 

1850:  Taenia  torulosa 

1861 :  Taenia  porulosa 

1884:  Taenia  torulosa 

1889:  Taenia  torulosa 

1891:  Taenia  torulosa 

1892:  Taenia  torulosa 

1894:  Ichthyotaenia  torulosa 

1896 :  Ichthyotaenia  torulosa 

1897 :  Ichthyotaenia  torulosa 

1901 :  Ichthyotaenia  torulosa 

1902:  Ichthyotaenia  torulosa 

1905 :  Ichthyotaenia  torulosa 

1905:  Proteocephalus  torulosus 

1909:  Ichthyotaenia  torulosa 

1911 :  Proteocephalus  torulosus 


Bloch 

1782 :11 

Batsch 

1786 :181-182 

Schrank 

1788 :49 

Frolich 

1791 :58-61 

Gmelin 

1790 :3081 

Viborg 

1795 :240 

Zeder 

1800:215,220 

Rudolphi 

1802a  :110-112 

Zeder 

1803 :352-353 

Rudolphi 

1810:111 

Rudolphi 

1819 :149-150 

Dujardin 

1845:584 

Diesing 

1850:514 

Van  Beneden 

1861 :162-163 

Zschokke 

1884:20,  in  part 

Lonnberg 

1889:15 

von   Linstow 

1891 :565 

Kramer 

1892 :55 

Lonnberg 

1894 :801-803 

Zschokke 

1896 :775 

von  Ratz 

1897 :159 

Fric  &  Vavra 

1901 :111-112 

Schneider 

1902 :24 

Schneider 

1905 :24-25 

Nufer 

1905 :75, 147,  in  ] 

Liihe 

1909:32 

La  Rue 

1911 :475 

119]  PROTEOCEPHALIDAE—LA  RUE  119 

Specific  Diagnosis:  Characters  of  the  genus.  Large  cestodes  65- 
600  mm.  long  by  1.2-2,25  mm.  broad.  Head  large,  prominent,  swollen, 
flattened  dorsoventrally,  without  rostellum,  without  fifth  sucker. 
Breadth  of  head  0.480-0.600  mm.,  thickness  about  0.300  mm.  Suckers 
circular,  prominent,  with  deep  cavities,  directed  anteriad  and  outward. 
Diameter  of  sucker  0.18-0.20  mm.  Neck  thick,  broad,  2-3  mm.  long, 
wrinkled.  First  proglottids  much  broader  than  long,  mature  and  ripe 
proglottids  almost  always  broader  than  long.  Maximum  length  of  ripe 
proglottids  1.0  mm.,  maximum  breadth  of  same  2.5  mm.  Proglottids 
fleshy,  well  delimited  by  deep  inter-segmental  furrows,  corners  of  seg- 
ments prominent.  End-proglottid  rounded  posteriorly.  In  its  posterior 
end  a  deep  indentation. 

Genital  pore  irregularly  alternating,  situated  near  middle  of  lateral 
proglottid  margin.  Vaginal  opening  dorsal  and  anterior  to  opening  of 
cirrus-pouch.  Cirrus-pouch  and  vagina  opening  into  a  common  genital 
sinus.  Testes  large,  0.16  by  0.08  mm.,  100-110  in  number.  Testes  in 
two  layers,  one  occupying  the  whole  dorsal  field  between  vitellaria, 
completely  covering  ovary.  Vas  deferens,  an  eccentric  mass  of  coils 
reaching  from  cirrus-pouch  to  middle  of  proglottid.  Cirrus-pouch 
about  0.255  mm.  long  by  0.085  mm.  broad,  extending  l^-%  across  pro- 
glottid breadth.  Cirrus  short,  not  heavy.  Ductus  ejaculatorius  describ- 
ing few  or  no  coils. 

Vaginal  opening  dorsal  and  anterior  to  cirrus-pouch.  Sphincter 
vaginae  very  weak,  situated  near  vaginal  opening.  Vagina  not  crossing 
cirrus-pouch.  Vitellaria  lateral,  voluminous,  follicles  large  and  closely 
packed  together.  Ovary  bilobed,  lobes  long,  thick,  and  irregular  in 
outline.  Ootype  and  oocapt  present.  Uterus  in  ripe  proglottids  with 
3-4  lateral  pouches  on  either  side.  Uterine  pores  not  observed.  Eggs 
provided  with  three  membranes.  Outer  membrane  hyaline,  0.055  mm., 
second  membrane  granular,  0.032  mm.,  embryo  about  0.021  mm.  in 
diameter. 


120  ILLINOIS  BIOLOGICAL  MONOGRAPHS 

Habitat :    In  intestine  of  host. 


[120 


Host 

Locality 

Collector 

Authority 

Idus  melanotus  (Cyprinus 

jeses)    (type  host) 
Idus  melanotus  (Cyprinus 

Batsch    (1786:181-182) 

jeses) 

Berlin 

Bloch 

Rudolphi   (1819:150) 

Idus  melanotus  {Cyprinus 

jeses) 

Greifswald 

Rudolphi 

Rudolphi   (1819:150) 

Idus  melanotus  (Cyprinus 

orfus) 
Leuciscus  leuciscus 

Frolich 

Rudolphi   (1819:150) 
Rudolphi   (1819:150) 

Zeder 

Leuciscus  leuciscus 

_„ 

von  Linstow  (1878:254) 

Leuciscus  leuciscus 

Podiebrad, 

Fric  and 

Fric  &  Vavra 

Bohemia 

Vavra 

(1901  :lii-ii2) 

Leuciscus  leuciscus 

Lake  Lucerne 

Xufer 

Nufer    (1905:75) 

Idus  melanotus  (Leuciscus  idus) 

Abildgaard 

Diesing  (1850:559) 
Diesing  (1850:514) 
Schneider    (1902:24 

Idus  melanotus  (Leuciscus  idus) 
Idus  melanotus  (Leuciscus  idus) 

(In  M.  C.  V.) 
Finland 

Schneider 

&  1905:24-25) 

Leuciscus  grislagine 

Sweden  (?) 

Tullberg 

Lonnberg   (1889:15) 

Albumus  bipunctatus 

(In  M.  C.  V.) 

Diesing 

Diesing  (1850:514) 

Alburnus  sp. 

Creplin 

Diesing  (1850:514) 

Albumus  lucidus 

von  Linstow    (1878:258) 

Alburnus  lucidus 

Lake  Geneva 

Zschokke 

Zschokke    (1884:20) 

Alburnus  lucidus 

Lake  Lucerne 

Kraemer 

Kraemer  (1892:55-71) 

Alburnus  lucidus 

v^on  Lin  stow 

von  Linstow   (1891:565) 

Alburnus  lucidus 

Zschokke 
Nufer 

Zschokke    (1896:775) 

Alburnus  lucidus 

Lake  Lucerne 

Nufer    (1905:75) 

Aspius  rapax 

Creplin 

Diesing    (1850:514) 

A  bra  mis  brama 

von  Linstow    (1878:258) 

Abramis  brama 

Lake  Balaton 

von  Ratz 

von  Ratz   (1897:159) 

Idus  melanotus 

von  Linstow  (1878:255) 

Pelecus  cultratus 

Lake  Balaton 

von  Ratz 

\on  Ratz   (1897:159) 

Gobio  fluviatilis 

Lake  Lucerne 

Xufer 

Nufer    (1905:75) 

Squalius  leuciscus 

Lake  Lucerne 

Nufer 

Nufer    (1905:75) 

Blicca  bjoerkna 

Lake  Lucerne 

Nufer 

Nufer    (1905:75) 

(  ?)Coregonus  fera 

Lake  Geneva 

Zschokke 

Zschokke  (1884:20) 

(?)Coregonus  exiguus  albellus 

Lake  Lucerne 

Nufer 

Nufer  (1905:75) 

(?)Coregonus  schinsii  helveiicus 

Lake  Lucerne 

Nufer 

Nufer  (1905:75) 

(?)Salmo  salvelinus 

Lake  Lucerne 

Nufer 

Nufer  (1905:75) 

(  })Perca  fluviatilis 

Luke  Lucerne 

Nufer 

Nufer  (1905:75) 

(?)Perca  fluviatilis 

Podiebrad, 

Fric  and 

Fric  &  Va\Ta 

Bohemia 

Vavra 

(1901 :iii-ii2) 

(?)Lota  vulgaris 

Lake  Geneva 

Zschokke 

Zschokke  (1884:20) 

121]  PROTEOCEPHALIDAE—LA  RUE  121 

Batsch  (1786:181-182)  first  described  this  species  from  Cyprinus 
jeses  giving  it  tlie  name  Taenia  torulosa.  His  description  is  here  quoted 
in  fuU: 

"Taenia  torulosa.    Der  rundgliedrige  Bandwurm. 

"Bloch   Preissschr.   S.  ii  nr.  4.     T.  2.  fig.  1-4.  10.  11.     Taenia  articulis  rotundis. 

Der  rundgliedrige  Bandwurm. 

Taenia   (loriformis)   capite  inermi,  obtuso,  osculis  per  paria  difformibus:  corpore 

toruloso,  articulis  orbicularibus  coUo  crenato,  elongata. 

"Diese  und  die  folgende  Art  haben  einen  abgestumpften,  die  nachsten  fiinf 
aber  einen  stumpfspitsigen  Scheitel,  bei  alien  sind  weder  die  Haken,  noch  der 
besondere  Russel  vorhanden  wie  bei  denen  vier  vorigen. 

"Der  rundgliedrige  Bandwurm  hat  rundliche  Glieder,  die  kurz,  dick,  und 
undurchsichtig,  und  wie  eine  Korallenschnur  an  einandergereiht  sind.  Am  Rande 
die  Miindungen. 

"Der  Hals  ist  kurz. 

"Der  Kopf  bewegt  seine  vier  Saugmiindungen  paarweis,  bald  in  halbmond- 
formige  Wiilste,  bald  in  die  Gestalt  eirunder  Oeffnungen.  Bisweilen  erweitert  er 
sie  alle  vier  in  eine  zirklrunde  Form,  und  alsdenn  verschwindet  ihr  sonst  sicht- 
barer  Ring. 

"Die  Eier  gehen  unter  dem  Presschieber  haufig  ab. 

"Die  Lange  betragt  zwei  Zoll,  die  Breite  eine  Linie. 

"Er  findet  sich  im  Aland  (Cyprinus  Jeses)  nebst  vielen  Nelkenbindwiirmem 
und  Kratzern." 

Batsch 's  drawings  of  the  head  are  reproduced  (Figs.  145,  146). 

Prior  to  Batsch  (1786)  one  other  investigator  reported  cestodes  of 
Cyprinidae  which  perhaps  may  be  referred  to  this  species.  Bloch 
(1782:11)  also  had  found  this  form  in  Cyprinus.  He  reported  it  under 
the  descriptive  name  Taenia  articulis  rotundis.  His  report  is  inaccessi- 
ble to  me.  Batsch  knew  of  Bloch 's  report  for  he  mentions  it.  Schrank 
(1788:49)  reported  this  species  under  the  name  Taenia  orbicularis  and 
he  mentioned  Taenia  articulis  rotundis  Bloch  as  a  synonym.  His  diag- 
nosis reads:     "Vier  Saugmiindungen  am  Kopfe;  die  Glieder  des  Kor- 

pers  tellerformig.    Taenia  orbicularis Wohnort,  im  Aland. ' ' 

Frolich  (1791:58-61)  described  a  cestode  from  Cyprinus  orfus  giving  it 
the  name  Taenia  simplex.  This  article  is  not  accessible  to  me.  Rudol- 
phi  (1802a  :110-112)  commented  on  Frolich 's  work  on  this  form  in  these 
words:  "Dass  er  (Frolich)  Blochs  Taenia  torulosa  nicht  citirt,  ist 
sehr  begreiflich,  da  Bloch  die  Glieder  ganz  anders  vorstellt.  Ich  glaube, 
dass  die  Taenia  simplex  nur  jiingere  Exemplare  derselben  Art  bezeich- 
nen."  Gmelin  (1790:3081)  gave  a  brief  diagnosis  of  Taenia  torulosa 
which  he  credited  to  Batsch  (1786)  tho  he  also  made  reference  to  Bloch 
(1782).  His  diagnosis  reads:  "T.  elongata  torulosa,  capite  obtuso,  collo 
crenato,  articulis  orbicularis :  osculis  geminis  difformis, Habitat 


122  ILUNOIS  BIOLOGICAL  MONOGRAPHS  [122 

in  Cyprino  Jese,  ad  2  pedes  longa,  capite  piano,  anierius  truncato,  coUo 
hrevi,  articulis  brevibtis  crassis." 

Viboi^  (1795)  reported  Taenia  cjrprini  idi  upon  which  Budolphi 
(1810:213)  commented  in  these  words:  "Taenia  Cyprini  Idi.  Viborg 
Ind.  Mus.  Vet.  Hafn.  p.  240.  n.  133.  T.  Cyprini  Idi. 

"Non  describitar,  forsan  tamen  ad  Taeniam  torulosam  (n.  22)  per- 
tinet,  quae  pluribus  in  Cyprinis  occurrit." 

Viboi^'s  writings  are  not  accessible  to  me.  Dr.  C.  W.  Stiles  in 
reply  to  a  letter  asking  for  information  on  this  species  wrote,  "Appar- 
ently Viborg  mentions  Taenia  idi  in  1795a  (See  author's  Catalogue),** 
referring  to  the  Index  Catalogue  of  Medical  and  Veterinary  Zoology. 
Zeder  (1800:215,  220)  reported  a  cestode  from  Cyprinus  under  the 
name  Rhyielminihus  cyprini  which  name  he  later  (1803:352-3)  consid- 
ered to  be  a  synonym  of  Halysis  torulosa.  His  last  diagnosis  reads: 
"HA LYSIS  TORULOSA.— fere  linearis,  plana;  capite  terete,  antice 
truncato,  cum  coUo  longo  continuo;  osculis  anticis  conicis;  corpore  cre- 
nato Habitat  in  intestinis  cyprinorum  jesis  et  leucisci." 

Rudolphi  (1802a  :110-12)  after  stating  a  brief  diagnosis  of  Taenia 
torulosa  presented  some  new  data  in  the  words  which  are  here  quoted: 

"Ich  habe  diesen  Wurm,  wie  Bloch,  im  Dannkanal  des  Alands,  Cyprinus 
Jeses,  im  April,  ausserst  haufig  angetroflfen,  aber  nie  iiber  einen  Fuss  lang,  da 
ihn  Bloch  hingegen  doppelt  so  gross  angetroffen  hat.  Der  Kopf  ist  von  Bloch  gut 
beschrieben  und  abgebildet,  die  Saugwarzen  und  der  ganz  Kopf  verandem  ihre 
Gestalt  alle  Augenblicke,  bald  stehen  ein  paar  Saugmiindiuigen  stark  hervor,  bald 
nur  eine,  bald  alle;  bald  erscheinen  sie  wie  tiefe  Napfschen,  bald  stehen  sie  hervor, 
bald  sieht  man  sie  zur  Halfte,  bald  ist  der  Rand  gleichsam  doppelt  u.  s.  w.  Uebri- 
gens  kann  ich  sie  schon  mit  blossen  Augen  sehen;  der  Kopf  ist  bald  mehr,  bald 
weniger  aufgeblasen,  und  immer  starker  als  der  Hals.  Der  Korper  ist  rundlich 
imd  wird  nach  hinten  allmalich  grosser;  dass  er  gegliedert  ist,  glaube  ich  gefunden 
zu  haben,  aber  so  deutlich  ist  er  es  nicht,  wie  man  es  auch  nach  der  Blochschen 
Zeichnung  (Taf.  II.  Fig.  i.)  glauben  sollte,  wo  man  in  einiger  Entfemung  vom 
Kopfe  schon  grosse  runde  Glieder  wahmimmt ;  bei  den  vielen  von  mir  unter- 
suchen  Exemplaren  nimmt  der  Korper  in  seinem  Verlauf  nur  schwach  zu,  und 
die  Glieder  sind  nur  schwach  unterschieden ;  auch  habe  ich  an  keinem  einzigen 
eine  SeitenoflFnung  bemerkt.  Diese  letztere  findet  man  aber  gewohnlich  nur  bei 
grossen  Exemplaren,  und  will  ich  also  darin  geme  Blochs  Beobachtungen  glauben." 

Rudolphi  (1810:111)  gave  a  diagnosis,  synonymy  and  description 
of  this  species  which,  because  it  admirably  sums  up  all  the  previous 
work  on  the  form  and  because  of  its  inaccessibility,  is  here  quoted 
verbatim : 

"Taenia  torulosa  Batsch. — Taenia:  capite  truncato,  osculis  orbicularibus  mar- 
ginatis,  collo  mediocri,  articulis  crassiusculis  (subquadratis)  subrotundis. 
Bloch  Abb.  p.  II.  Tab.  2.  fig.  1-4.    Taenia  articulis  rottmdis. 


123]  PROTEOCEPHALIDAE—LA  RUE  123 

Batsch  Bandw.  p.  i8i  n.  27.  fig.  105-108.    Taenia  torubsa. 

Schrank  Verz.  p.  49.  11,150.    T.  orbicularis. 

Gmel.  Syst.  n.  p.  3081.  n.  85.    T.  torulosa. 

Frolich  im  Naturf.  25.  p.  58-61.  T.  3.  fig.  4-6  T.  simplex. 

Zeder  Nachtrag  p.  220.    Rhytelminthus  Cyprini. 

Rudolphi  in  Wied.  Arch.  iii.  i.  p.  no.    Taenia  torulosa. 

Zeder  Naturg.  p.  352.  n.  39.     Halysis  torulosa. 

"Hab.  in  ventriculo  vel  intestinis  Cyprinorum.  In  Cyprino  lese  Bloch  et  ego 
Aprili,  in  Orfa  Frolichius  eodem,  in  Leticisco  Zederus  Majo  mense,  reperimus. 

"Descr.  Vermes  Zederiani  septem  ad  novem  lineas  longi,  dimidiam  lati ; 
Frolichiani  duos  ad  quinque  pollices  longi,  postice  lineam  lati ;  mei  pedem  non 
superantes  vix  lineam  dimidiam  lati ;  Blochiani  duos  pedes  longi,  lineam  et  quod 
excurrit  lati. 

"Caput  truncatum,  depressum,  sub  motu  polymorphum,  inflatum  et  osculorum 
directionem  sum  mopere  mutans ;  haec  circularia,  concava,  marginata,  margine 
simplici  \el  duplici,  interdum  occultato ;  mox  omnia  quatuor,  mox  duo  tantum  in 
conspectum  veniunt,  ceterum  oculis  nudis  usurpanda.  Collum  depressum,  medio- 
cre, a  capite  crassiore  discretum.  Corpus  crassiusculum,  subaequale,  margine  cre- 
natum,  articulis  confluentibus,  margine  postico  vix  incumbentibus. 

"Obs.  I.  Blochius  specimina  sex  maxima  vidit,  eorumque  articulos  quam  in 
meis  magis  discretos,  ovis  farctos,  et  foramina  marginalia  (haec  tamen  non  ulte- 
rius  exposita,  neque  delineata)  observavit.  Frolichiani  vermes  a  meis  vix  ulla 
nota  differunt,  nisi  quod  collum  longum  dicat,  quod  ipse  mediocre,  Blochius  breve 
vocaverit,  hoc  autem  vermis  totius  ratione  habita  facile  explicatur ;  in  vermlbus 
enim  bipedulibus  collum  vix  pollicare  Blochio  breve  visum,  quod  Frolichio  in 
bivel  quinquepollicaribus  longum,  mihi  in  pedalibus  mediocre  fuerit. 

"Obs.  2.  Zederiani  vermes  valde  pusilli  fuere,  ideoque  a  nostria  megis  dis- 
tant; ab  osculis  quatuor  suctoriis  totidem  canales  oriebantur,  in  corpore  progressi, 
tandem  in  unicum  vas  abeuntes,  quod  in  cauda  obtuso-acuta  terminabatur.  Haec 
vasa  mihi  non  visa,  sed  vermes  illi  tenelli  plani,  ideoque  pellucidi  fuere,  nostri 
muto  crassiores,  fere  teretiusculi.  Zederus  etiam  postea  ipse  Rhytelminthum  cy- 
prini olim  sibi  dictum  ad  T.  torulosum  pertinere  suspicatus  est,  quod  olim  in 
diario  Wiedemanniano  aeque  indicaveram.  Conf.  etiam  Taeniam  Cyprini  Idi  n. 
114.  dictam." 

Rudolphi  (1819:150)  stated  that  this  species  had  been  found  by 
Bloch,  Frolich,  Zeder  and  himself  in  certain  species  of  the  Cyprinidae. 
Dujardin  (1845:584)  made  no  record  of  finding  this  species.  He  col- 
lected his  data  from  descriptions  by  Zeder,  Frolich,  and  Bloch,  His 
description  reads:  '* — Long  de  16  a  20  mm,  et  large  de  1,12  mm. 
(Zeder),  ou  large  de  50  a  135  mm,  et  large  de  225  mm  (Froelich),  ou 
long  de  330  mm,  large  de  1.2  mm,  (Rud.),  ou  enfin  long  de  660  mm, 
large  de  225  a  (?)  (Bloch)  ; — tete  tronquee,  de  forme  tres- variable, 
ainsi  que  ses  ventouses  dont  le  bord  est  saillant; — trompe  nuUe; — cou 
de  longueur  mediocre ; — articles  assez  epais,  presque  ronds. ' ' 


124  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [124 

Diesing  (1850:514)  found  this  species  in  Leuciscus  idus  and  in 
Alhurnus  bipunctatus  and  further  stated  that  Creplin  found  it  in  As- 
pius  rapax  and  in  Alburnus  sp.  Diesing 's  diagnosis  added  a  little  to 
the  previous  knowledge  of  the  form.  His  diagnosis  reads:  ^^ Caput 
latum  depressum  truncatum,  acetabulis  lateralibus  limbo  prominulo. 
Collum  longum.  Articuli  teretiusculi  subaequales  longi  confluentes. 
Aperturae  genitalium.  .  .  Longit.  7-10";  latit.  i/^-l'"."  Van  Bene- 
den  (1861:162-163)  reported  finding  this  form  in  several  species  of 
Cyprinidae  ("plusieurs  especes  de  cyprins").  Altho  he  gave  almost 
no  descriptive  data  a  comparison  of  his  figures  with  those  of  Batsch 
leaves  little  doubt  that  his  Taenia  porulosa  is  identical  and  synonymous 
with  T.  torulosa  Batsch.  Porulosa  probably  is  a  misspelling  of  torulosa. 
Some  authors  have  denied  the  synonymy  of  these  names  but  their  con- 
tention has  no  support  of  facts.  Van  Beneden  failed  to  state  the  locality 
of  his  catch.  Von  Linstow  (1878  and  1889)  catalogued  the  hosts  from 
which  this  species  had  been  collected.  He  added  nothing  to  the  descrip- 
tion of  the  species. 

Zschokke  (1884:20)  stated  that  he  found  Taenia  torulosa  in  Core- 
gonus  fera,  Lota  vulgaris  and  Alhurnus  lucidus  from  Lake  Geneva. 
Since  Zschokke  reported  Taenia  longicollis  and  T.  ocellata  from  Core- 
gonus  fera  in  addition  to  Taenia  torulosa  it  is  highly  probable  that  all 
of  his  specimens  from  Coregonus  belonged  to  one  of  the  last  named  ces- 
tode  species  rather  than  to  T.  torulosa.  Zschokke  further  states  that 
his  specimens  were  too  young  to  permit  him  to  recognize  any  trace  of 
the  internal  organs.  These  statements  as  to  a  probable  misdetermina- 
tion  of  species  apply  equally  well  to  his  parasite  report  on  Lota  vulgaris 
in  which  case  he  found  Taenia  ocellata  in  addition  to  Taenia  torulosa. 
His  four  young  specimens  from  Alhurnus  lucidus  may  have  been  Taenia 
torulosa  if  only  the  question  of  host  be  considered.  Of  these  four  speci- 
mens he  wrote:  '*La  tete  etait  large,  tronquee,  les  ventouses  tres 
fortes  et  saillantes.  Le  cou  etait  de  longeur,  mediocre,  la  segmentation 
en  articles  peu  accusee."  Lonnberg  (1889:15)  reported  the  finding  of 
Taenia  torulosa  by  Professor  Tullberg.  He  gave  no  description  and 
did  not  state  the  locality  of  the  collection.  Von  Linstow  (1891:565) 
found  Taenia  torulosa  in  Alhurnus  lucidus.    He  gave  no  data. 

Kramer  (1892:55)  found  Taenia  torulosa  in  Alhurnus  lucidus, 
Lake  Lucerne.  He  examined  more  than  150  specimens  of  Coregonus 
fera  without  finding  a  single  specimen  of  Taenia  torulosa,  and  among 
numerous  specimens  of  Alhurnus  lucidus  but  a  single  host  was  infected 
with  Taenia  torulosa.  He  made  the  first  careful  morphological  and 
histological  study  of  this  species.  Lonnberg  (1894:801-803)  included 
Taenia  torulosa  in  a  list  of  species  of  his  new  genus  Ichthyotaenia. 


125]  PROTEOCEPHALIDAE—LA  RUE  125 

Zschokke  (1896:775)  found  Ichthyotaenia  torulosa  (Batsch)  in  the  in- 
testine of  Leuciscus  leuciscus  L.  He  found  it  in  no  other  hosts.  Von 
Ratz  (1897:159)  reported  finding  Ichthyotaenia  torulosa  in  Abramis 
hrama  L.  and  in  Pelecus  cultratus  L.  in  Lake  Balaton,  Hungary,  but 
he  gave  no  description.  Fric  and  Vavra  (1901:111-112)  reported  what 
they  considered  to  be  Ichthyotaenia  torrulosa  (Batsch)  (misspelling 
for  torulosa)  from  Leuciscus  leuciscus  and  Perca  fluviatUis,  Podiebrad, 
Bohemia.  The  description  of  their  specimens  agrees  very  well  with 
that  of  Proteocephalus  percae  and  a  part  or  all  of  their  specimens  may 
belong  to  that  species.  The  Hasling  (Leuciscus  leuciscus)  would  be  a 
new  host  for  P.  percae  and  it  may  be  that  their  specimens  from  that 
host  are  as  they  identified  them,  i.  e.,  P.  torulosus.  Moreover,  it  is  not 
impossible  that  Fric  and  Vavra  failed  to  distinguish  between  the  two 
species  of  cestode,  P.  percae  and  P.  torulosus.  It  is  impossible  to  make 
a  complete  determination  from  their  data.  Schneider  (1902:24)  found 
Ichthyotaenia  torulosa  in  Leuciscus  idus  in  Finland.  Three  years  later 
he  (1905:24-25)  briefly  but  concisely  described  this  species  from  Leu- 
ciscus idus.  His  description  agrees  very  well  with  that  of  Kramer 
(1892). 

Nufer  (1905:75)  reported  Proteocephalus  torulosus  (Batsch)  from 
Perca  fluviatilis,  Alburnus  lucidus,  Squalius  leuciscus,  Blicca  hjoerkna, 
Oohio  fluviatilis,  Coregonus  exiguus  aVhellus,  C.  schinzii  helveticus, 
Salmo  salvelinus.  Some  of  Nufer 's  report  is  open  to  doubt.  In  59 
specimens  of  Perca  fluviatilis  he  not  only  failed  to  find  the  species  which 
Zschokke  found  in  that  host  but  he  found  P.  longicollis  and  P.  torulo- 
sus, a  species  which  only  once  before  (Fric  and  Vavra,  1901)  had  been 
reported  from  that  host.  In  that  instance  attention  has  been  called  to 
the  fact  that  the  specimens  in  question  were  probably  P.  percae  and  not 
P.  torulosus.  Nufer 's  P.  torulosus  and  his  P.  longicollis  may  have  been 
P.  percae.  Nufer 's  next  four  hosts  after  Perca  fluviatilis  are  all  Cy- 
prinidae,  from  which  group  this  species  has  heretofore  been  reported. 
It  is  extremely  doubtful  if  the  Salmonidae  harbor  this  species.  Zschok- 
ke, who  in  1884  reported  young  specimens  of  Taenia  torulosa  from 
Coregonus  fera,  in  1896  found  this  parasite  only  in  Alhurnus  lucidus. 
Zschokke 's  paper  (1896)  contained  the  results  of  his  investigations  on 
more  than  1600  fish  from  Lake  Geneva  and  the  Rhine  hence  his  data 
were  fairly  comprehensive.  Moreover,  from  the  three  species  of  Salmon- 
idae in  which  he  claimed  to  have  found  P.  torulosus  Nufer  reported 
two  other  species  of  Proteocephalus,  namely,  P.  longicollis  and  P.  ocel- 
latus.  Here  again  Nufer  probably  made  a  misdetermination.  The 
writer  has  shown  that  Nufer 's  work  (see  discussion  of  P.  macrocepha- 
lus)   is  untrustworthy  and  in  the  determination  of  this  species  there 


126  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [126 

seems  to  be  no  exception.  That  Nufer  made  a  misdetermination  of  his 
P.  longicollis  is  not  at  all  improbable  since  specimens  labelled  ^'Taenia 
longicollis  aus  Forelle"  which  have  been  received  by  Professor  Ward 
from  Professor  Zschokke  have  proved  to  belong  to  a  different  species. 
Nufer 's  tabulated  description  of  P.  longicollis  (Nufer  1905:147)  does 
not  agree  in  some  important  points  with  von  Linstow's  description  yet 
it  is  very  manifest  that  he  drew  on  von  Linstow's  description  for  a 
large  part  of  his  data.  Likewise  parts  of  Nufer 's  description  of  P. 
torulosus  (Nufer  1905:147)  fail  to  agree  with  the  descriptions  of  that 
species  by  Schneider  and  Kraemer.  Liihe  (1909:32)  gave  a  short  spe- 
cific description  of  this  form  under  the  name  of  Ichthyotaenia  torulosa 
(Batsch).  La  Rue  (1911:475)  listed  this  form  among  other  species  of 
Proteocephalus. 

The  following  study  is  based  upon  some  of  Schneider's  specimens 
which  Prof.  H.  B.  Ward  secured  from  Professor  Levander  at  Helsing- 
fors.  This  material  is  labelled  ^^Ichthyotaenia  torulosa  Batsch.  Leucis- 
cus  idus.  Porkala.  Juni  1901.  det.  G.  Schneider."  It  now  bears  the 
number  10.121  in  Professor  Ward's  collection.  From  this  material 
frontal  and  transverse  sections  have  been  made  and  some  heads  have 
been  studied  in  glycerine.  A  careful  comparison  of  Schneider's  (1905) 
and  Kraemer 's  (1892)  descriptions  shows  very  good  agreement.  Zschok- 
ke's  (1884)  description  in  parts  agrees  pretty  well  with  these  two.  Liihe 
(1909)  seems  to  have  based  his  specific  description  largely  on  the  work 
of  Zschokke,  Kramer,  and  Schneider.  Data  from  these  sources  are  used 
in  this  description. 

This  is  one  of  the  larger  species  of  Proteocephalus  infesting  fish. 
In  length  it  varies  from  65  to  600  mm.  Specimens  of  the  latter  length 
are  rare  and  have  been  reported  only  by  some  of  the  earlier  investigators. 
The  longest  complete  strobila  examined  by  the  writer  was  110  mm.  long. 
The  breadth  of  the  strobila  varies  from  1.2-2.25  mm.  The  head  (Fig.  7) 
is  large  and  very  prominent,  somewhat  swollen  in  appearance,  flattened 
dorso-ventrally,  and  somewhat  flattened  at  the  apex.  Schneider  found 
it  0.500  mm.  broad.  The  heads  in  the  writer's  material  measured  0.450- 
0.600  mm.  broad  by  about  0.320  mm.  thick.  The  suckers  are  very  prom- 
inent, nearly  circular  in  outline,  with  deep  cavities  and  strong  muscu- 
lature. They  are  directed  anteriad  and  a  little  outward.  The  suckers 
are  0.200  mm.  in  diameter  according  to  Schneider  and  about  0.180-0.200 
mm.  in  the  heads  examined  by  me.  A  fifth  sucker  is  not  present.  In 
sections  of  the  head  the  writer  was  unable  to  find  even  a  trace  of  a 
vestigial  fifth  sucker.  Nufer  (1905:147)  stated  that  the  head  of  a  37 
mm.  specimen  measured  0.255  mm.  broad  and  the  suckers  of  the  same 
were  0.10  mm.  in  diameter.    Nufer 's  measurements  of  the  head  do  not 


127]  PROTEOCEPHALIDAE—LA  RUE  127 

agree  at  all  with  the  measurements  made  by  Schneider  or  the  writer. 
Neither  Kramer  nor  Zsehokke  give  measurements  of  the  head  and  suck- 
ers but  they  state  that  the  head  is  large,  the  suckers  large  and  round. 
Nufer's  specimens  which  infested  Perca  fluviatilis  most  certainly  could 
not  have  been  P.  torulosus  (Batsch)  but  were  either  P.  percae,  P.  fallax 
La  Rue,  or  an  undescribed  species. 

Batsch  drew  the  head  of  P.  torulosus  as  a  broad  structure  with  large 
and  prominent  suckers.  His  figures  which  have  been  reproduced  (Figs. 
145,  146)  should  be  compared  with  the  drawing  of  a  head  (Fig.  7) 
derived  from  material  sent  to  Professor  Ward  by  Professor'  Levander. 
There  is  remarkable  agreement  in  form  of  head  and  of  suckers.  The 
thick  neck  is  about  2-3  mm.  long  by  about  0.20-0.30  mm.  broad.  It  is  much 
wrinkled  thus  making  difficult  the  determination  of  the  length.  It 
passes  over  gradually  into  the  first  proglottids  which  are  much  broader 
than  long.  Mature  and  ripe  proglottids  are  almost  always  broader 
than  long.  The  maximum  length  observed  was  1.0  mm.  and  the  maximum 
breadth  2.5  mm.  Many  nearly  ripe  proglottids  measure  about  0.29 
mm.  long  by  1.30  mm.  broad.  The  proglottids  are  thick  and  fleshy  with 
well  defined  limits.  Indentations  between  the  proglottids  are  deep  and 
the  corners  of  the  latter  are  quite  pronounced.  The  end  proglottid  is 
rounded  posteriorly.  At  its  posterior  end  it  has  a  deep  indentation 
into  which  the  excretory  ducts  discharge. 

According  to  Kramer  the  excretory  ducts  in  the  head  and  neck 
region  are  very  prominent  with  many  anastomosing  coils.  In  the  neck 
region  he  found  many  branches  of  the  excretory  vessels  leading  to  the 
exterior.    Sections  made  by  the  writer  failed  to  show  these  relations. 

The  genital  openings  are  irregularly  alternating,  and  are  situated 
near  the  middle  of  the  margin  of  the  segment.  The  vagina  opens  not 
strictly  anterior  to  the  cirrus-pouch  but  somewhat  dorsal  thereto.  Kra- 
mer incorrectly  states  that  the  vagina  and  cirrus-pouch  open  near  each 
other  but  not  into  a  common  atrium.  The  testes  are  large,  spheroidal 
or  by  pressure  they  may  be  sometimes  rendered  polyhedral.  They 
measure  according  to  Schneider  about  0.160  mm.  in  sagittal  by  0.080 
mm.  in  frontal  diameter.  My  measurements  are  about  the  same  as 
these.  The  testes  lie  in  two  layers,  the  dorsal  layer  (Fig.  78)  covering 
the  entire  area  bounded  by  the  vitellaria  and  the  anterior  and  pos- 
terior margins  of  the  proglottid.  Thus  the  ovary  is  completely  covered 
by  the  dorsal  layer  of  testes  about  70  in  number.  In  the  more  ventral 
layer  there  are  about  30-40  testes.  Hence  the  total  number  of  testes  is 
about  100-110.  Kramer's  drawing  (Fig.  184)  shows  a  small  number 
of  testes  but  since  it  shows  the  ventral  view  of  the  worm  it  cannot  show 
a  large  number.    The  coils  of  the  vas  deferens  form  a  thick  mass  ex- 


128  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [128 

tending  from  the  cirrus-pouch  to  the  mid-field  of  the  segment.  Kramer 
(Fig.  184)  figures  the  coils  of  vas  deferens  lying  in  the  middle  of  the 
segment.  The  cirrus-pouch  is  short  and  relatively  thick.  Its  length  is 
about  0.255  mm.  and  its  breadth  0.085  mm.  The  ratio  of  its  length  to 
the  proglottid  breadth  is  about  1 A  according  to  G.  Schneider,  1 :3  in 
Kraemer's  drawing,  and  1:4-1:6  in  my  preparations.  The  cirrus  is 
short  and  not  very  thick.  Kraemer's  description  of  the  cirrus  is  faulty 
in  that  he  said  that  the  cirrus  was  armed  with  recurved  hooks  which 
extend  back  into  the  tissue  of  the  cirrus  itself.  There  are  no  hooks. 
The  protruded  cirrus  has  not  been  seen  by  the  writer.  The  ductus 
ejaculatorius  forms  but  a  very  few  coils  or  instead  of  coils  it  may  lie 
in  sinuous  curves  within  the  cirrus-pouch. 

The  vagina  which  opens  mostly  dorsal  to  the  cirrus-pouch  has  a 
very  weak  sphincter  muscle  situated  near  the  opening  into  the  atrium. 
This  sphincter  vaginae  is  made  up  of  a  few  strands  of  circular  muscle 
fibers  which  may  be  easily  overlooked.  In  its  course  to  the  interovarial 
space  the  vagina  does  not  cross  the  cirrus-pouch  but  passes  dorsal  and 
anterior  to  the  pouch.  A  small  receptaculum  seminis  is  present  just 
anterior  to  the  ovary.  The  vitellaria  are  lateral,  voluminous,  and  fol- 
licular. The  follicles  are  large  and  closely  packed  together.  The  bi- 
lobed  ovary  is  large,  weU  developed,  thick  and  irregular  in  outline,  but 
the  lobes  are  not  as  slender  as  they  are  shown  in  Kraemer's  figure 
(compare  Figs.  79  and  184).  In  proglottids  1.3  mm.  broad  the  ovary 
may  have  a  span  of  0.80  mm.  and  the  lobes  may  be  0.130-0.140  mm. 
thick.  A  muscular  oocapt  and  an  ootype  are  present.  The  uterus  in 
ripe  proglottids  has  3-4  lateral  out-pocketings  on  either  side.  The  ute- 
rine pores  have  not  been  observed.  The  eggs  were  not  described  by 
Kramer  and  Schneider.  The  outer  membrane  is  thin  and  hyaline, 
0.055  mm.  in  diameter.  The  granular  second  membrane  has  a  diameter 
of  about  0.032  mm.  and  the  embryo  about  0.021  mm.  A  delicate  mem- 
brane closely  invests  the  embryo. 

P.  torulosus  differs  from  many  other  species  of  the  genus  by  its 
large  size,  and  from  a  large  number  of  the  species  through  its  lack  of  a 
fifth  sucker.  In  maximum  length  and  breadth  of  its  strobila  it  is  the 
largest  species  of  Proteocephalus  yet  described.  In  the  width  of  the 
head  and  diameter  of  the  suckers  it  is  exceeded  only  by  P.  amhloplitis 
and  P.  perplfixus.  In  observed  length  P.  torulosus  greatly  exceeds  these 
species — and  it  may  be  differentiated  from  them  by  its  lesser  number 
of  uterine  pouches,  its  weaker  sphincter  vaginae,  its  shorter  cirrus- 
pouch,  its  double  layer  of  testes,  and  by  the  arrangement  of  the  dorsal 
layer  of  the  same.  It  is  greatly  different  from  the  other  North  Ameri- 
can forms  thus  far  described.    Among  the  European  forms  P.  torulosus 


129]  PROTEOCEPHALIDAE—LA  RUE  129 

is  approached  in  size  only  by  P.  longicollis.  The  latter,  however,  has 
a  well  developed  fifth  sucker  and,  according  to  von  Linstow,  a  much 
smaller  number  of  testes.  P.  torulosus  may  be  differentiated  from  P. 
percae  and  P.  cernuae  by  its  larger  head,  larger  suckers,  lack  of  a  fifth 
sucker,  more  numerous  testes  and  the  different  arrangement  of  the  same, 
and  by  its  smaller  number  of  uterine  pouches.  P.  torulosus  is  much 
larger  than  P.  macroccphalus.  It  hds  a  larger  head,  larger  suckers, 
fewer  uterine  pouches  and  a  different  arrangement  of  the  dorsal  layer 
of  testes  than  the  latter  species.  P.  torulosus  is  so  different  from  the 
other  described  species  of  Proteocephalus  that  a  danger  of  its  being 
confused  with  them  is  scarcely  possible. 

'i 
PROTEOCEPHALUS  MACROCEPHALUS  (Creplin) 

[Figs.  1,  47-49,  171] 


1825: 

Taenia  macrocephala 

Creplin 

1825 

1845: 

Taenia  macrocephala 

Dujardin 

1845:585 

1850: 

Taenia  macrocephala 

Diesing 

1850 :513-514 

1859: 

Taenia  macrocephala 

Molin 

1859 :13 

1875: 

Taenia  macrocephala 

von  Linstow 

1875 

1889: 

Taenia  dilatata 

Linton 

1889 :488-489 

1893: 

Taenia  macrocephala 

Olsson 

1893 

1896: 

Ichthyotaenia  macrocephala 

Riggenbach 

1896 :267 

1897: 

Taenia  dilatata 

Linton 

1897:425 

1897: 

Taenia  macrocephala 

Stossich 

1897 :7 

1898: 

Taenia  macrocephala 

Stossich 

1898a  :113-114 

1898: 

Taenia  macrocephala 

Miihling 

1898 :37,  69 

1901: 

Taenia  dilatata 

Linton 

1901 :435 

1903: 

Ichthyotaenia  hemisphaerica 

Schneider 

1903 :29 

1905: 

Ichthyotaenia  macrocephala 

Schneidei 

1905 :10, 17-19 

1905: 

Proteocephalus  macroccphalus 

Nufer 

1905 :134-152 

1909: 

Ichthyotaenia  macrocephala 

Liihe 

1909:33-34 

1911: 

Proteocephalus  macroccphalus 

La  Rue 

1911 .475 

Specific  Diagnosis:  The  characters  of  the  genus.  Adult  cestodes 
possessing  strobila  as  much  as  40  cm.  long  by  1-1.8  mm.  broad  or  per- 
haps more.  Segmentation  quite  distinct.  Proglottids  very  numerous; 
first  much  broader  than  long;  mature,  broader  than  long  or  nearly 
quadrate ;  ripe,  longer  than  broad.  Neck  5.0-7.0  mm.  long,  0.1-0.25  mm. 
broad.  Head  globose,  flattened  dorsoventrally,  0.28-0.30-0.32  mm.  broad, 
0.15-0.16  mm.  thick.     Suckers    0.095-0.106    mm.  in    diameter.     Fifth 


130 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


[130 


sucker  vestigial  and  deeply  set  in  the  tissue  of  the  head,  0.025  mm.  in 
diameter.  Sexual  organs  as  in  genus.  Genital  aperture  marginal,  near 
middle  of  proglottid.  Vagina  anterior  and  vaginal  opening  dorsal  to 
cirrus-pouch.  Testes  100-120,  irregularly  scattered  between  vitellaria, 
lying  in  one  or  two  layers,  usually  one.  Cirrus-pouch  short,  about  0.16 
mm.  long.  Ratio  of  length  of  cirrus-pouch  to  proglottid  breadth  1:6- 
1:8.  Uterine  pouches  7-11-12-14  on  either  side.  Embryos  0.017-0.019- 
0.021  mm.  in  diameter. 

Habitat:     In  intestine  of  Anguilla  vulgaris   (type  host),  and  A. 
chrysypa  Raf. 


Host 

Locality 

Collector 

Anguilla  vulgaris 

Greifswald 

(type  locality) 

Creplin 

u                       « 

Rennes 

Dujardin 

«                    <( 

Padua 

Molin 

«                    « 

von  Linstow 

«                    <( 

<<                    « 

Narenta 

Stossich 

«                  « 

Trieste 

Stossich 

«                  « 

Memel, 

E.    Prussia 

Miihling 

«                    « 

Finland 

Schneider,  G. 

«                    « 

Lake  Lucerne 

Nufer 

"  chrysypa  Raf. 

Wood's  Hole,  Mass. 

Linton 

«           « 

((           «  ,       « 

Linton 

«           « 

Sebago  Lake,  Me, 

H.  B.  Ward 

Authority 


Creplin    (1825:69-71) 
Dujardin    (1845:585) 
Molin   (1859:13) 
von   Linstow    (1875) 
Olsson    (1893) 
Stossich   (1897:7) 
Stossich   (18983:113-114) 

Muhling  (1898:37-69) 
Schneider,  G.  (1903:29) 
Nufer  (1905:134-152) 
Linton  (1889:488-489) 
Linton  (1897:425-426) 
LaRue  (the  present  paper) 


Creplin  (1825)  found  this  species  in  the  intestine  of  Anguilla  vul- 
garis at  Greifswald.    His  diagnosis  and  description  are  here  given: 

"Taenia  macrocephala:  T.  capite  elongato,  antice  crassiore,  osculis  globosis 
anticis,  rostello  brevissimo  obtuso,  collo  brevi,  articulis  anterioribus  brevibus  ob- 
tusis,  sequentibus  subquadratis,  horum  lemniscis  marginalibus  vage  alternis. 

Hab.  In  intest.  Maraenae  Anguillae  Martio,  April,  Majo  et  Junio  banc 
Taeniam  reperi  aut  singulum  aut  minima  tamen  copia. 

Descr.  Vermes  aliquot  lineas  usque  ad  octo  poUices  longi,  majores  medio 
corpore  lineam  cum  quarta  vel  dimidia  ejus  parte  lati,  subdiaphani,  albissimi. 

Caput  oblongum,  antice  latius,  rostello  brevissimo,  obtuso  instructum,  inerme, 
et  quatuor  osculis  (quorum  orificia  non  vidi)  globosis,  prominentibus,  anticis, 
instructum.  Collum  breve,  rugosum,  cum  capite  continuum.  Articuli  antici  bre- 
vissimi,  inaequales,  obtusi,  passim  medio  constricti,  sequentes  sensim  latitudine  et 
longitudine  crescunt,  maximam  partem  quidem  latiores,  quam  longi  manent,  de- 


131]  PROTEOCEPHALIDAE—LA  RUE  131 

mum  vero  quadrat!  et  ultimi  adeo  paulo  longiores,  quam  lati  evadunt.  Omnes 
per  articulos  decurrit  ad  margines  laterales  utrinque  linea  alba  opaca.  Foramina 
articulorum  posticorum  marginalia  vage  alterna,  ex  quibus  cirrhus  propendet  bre- 
vis,   subtilissimus. 

Obs.  I.  Reperi  primo  hujus  vermis  specimen  unicum  valde  parvum  d.  2y 
Mart.  1822,  postea  d.  23.  Apr.  ej.  anni  iterum  parva  specimina  una  cum  Bothrio- 
cephalis  clavaecipitibus  itidem  parvis  demum  vero  specimen  octopoUicare  d.  23. 
Maji  1822,  et  secundum  specimen  magnum  d.  5.  Junii  1823,  simul  cum  illi  adjacente 
Bothr.  clavaecipite  mediocre.  Vi\X)s  examinae  microscopii  ope  non  mihi  licuit,  ut 
majorem  descriptionis  partem  de  verme  in  spiritu  servato  dare  debuerim. 

Obs.  2.  Specimen  Junio  inventum  vivacissimum,  aquae  frigidae  immissum, 
valdopere  se  movebat  et  contorquebat,  bisque  motibus  maximam  ovorum  copiam 
demittebat  subglobosorum,  ut  tota  aqua  inde  turbida  fieret. 

Obs.  3.  Hujus  vermis  caput  sine  dubio  O.  F.  Miiller  (Vid.  Schriften  Berl. 
Naturf.  I.  p.  208.  cit.  in  Rudolph.  Entozoologiae  T.  II.  P.  2.  p.  39.)  vidit,  cum 
Bothriocephalo  clavaecipiti  (Taeniae  Anguillae  sibi  dicto,  a  nostra  taenia  omnibus 
notis  diversissimo)   caput  osculis  quattuor  instructum  adscriberet." 

Dujardin  (1845:585)  reported  and  described  specimens  of  this  spe- 
cies in  these  words: 

"Tenia  de  I'anguille.  Taenia  macrocephala. — Long  de  8  a  220  mm,  large  de 
(?)  a  -Ti-Zy  mm; — tete  allongee,  plus  epaisse  en  avant,  avec  une  trompe  tres-courte, 
obtuse ; — ventouses  globuleuses,  dirigees  en  avant ; — cou  court ; — premiers  articles 
courts,  obtus,  les  suivant  presque  carres,  avec  les  orifices  genitaux  irreguliere- 
ment  alternes,  et  les  penis  courts,  tres-minces,  pendants. 

"M.  Creplin  la  trouve  plusieurs  fois,  a  Greifswald,  dans  I'intestin  de  I'an- 
guille. — J'ai  trouve  aussi  a  Rennes,  dans  une  anguille,  trois  jeunes  tenias  longs 
de  6  mm  et  10  mm  et  13.5  mm,  larges  de  0.25  mm  a  0.3  mm;  ayant  la  tete  large 
de  0.33  mm  a  048  mm,  sans  trompe,  et  les  ventouses  larges  de  0.106,  dirigees  en 
avant," 

Diesing  (1850:513-514)  added  nothing  to  the  earlier  descriptions. 
Molin  (1859)  reported  a  specimen  of  this  species  taken  from  AnguUla 
vulgaris  at  Padua  in  March,  1859.  He  did  not  describe  it.  Von  Lin- 
stow  (1875)  very  insufficiently  described  P.  macrocephalus  from  A71- 
guilla  vulgaris  in  the  following  words  (The  quotation  is  taken  from 
Nufer  1905 :135)  : 

"Der  Skolex  ist  gegen  den  folgenden  Korper  nicht  abgesetzt;  die  vier  grossen 
Saugnapfe  haben  einen  Durchmesser  von  0.166  mm.;  dazwischen  steht  ein  viel 
kleinerer,  funfter,  scheitelstandiger,  von  0.026  mm.  Durchmesser,  der  oft  schwer 
aufzufinden  ist.  Die  Cirren  sind  0.2  mm.  lang  und  0.06  mm.  breit ;  am  ausseren 
Drittel  zeigen  sie  eine  Einschniirung,  sodass  ungefahr  die  Gestalt  der  Kegel  ent- 
steht,  wie  sie  beim  Kegelschieben  iiblich  sind ;  sie  ragen  nur  wenig  mit  der 
Spitze  iiber  den  Rand  des  Gliedes  hinaus  und  stehen  unregelmassig  abwechselnd. 


132  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [132 

Die  aussere  EihuUe  ist  hyalin  und  umgibt  das  Ei  weitlaufig,  denn  sie  hat  einen 
Durchmesser  von  0.089  mm.,  wahrend  der  der  innern  Eihiille  nur  0.029  mm-  und 
der  der  Eizelle  selber  0.023  mm.  betragt.  Die  Embryonalhakchen  sind  sehr  fein; 
sie  haben  eine  Lange  von  0.006  mm." 

Von  Linstow's  measurements  of  the  suckers  much  exceed  those  given 
by  Schneider  and  the  writer  for  this  species.  He  also  reported  a  small 
fifth  sucker  that  was  very  difficult  to  see.  The  writer  is  inclined  to  the 
belief  that  this  was  the  small  vestigial  fifth  sucker  which  in  some  Pro- 
teocephalids  may  be  seen  in  favorable  toto  preparations. 

Linton  (1889:488-489)  described  some  cestodes  from  Anguilla 
chrysypa  Raf.,  caught  on  our  eastern  coast  as  a  new  species,  Taenia 
dilatata  in  these  words: 

"Head  small,  truncate,  or,  in  living  specimens,  slightly  prominent  in  front 
Acetabula  nearly  circular,  directed  a  little  forward.  Neck  rugose,  very  long, 
very  contractile  and  dilatable,  narrow  in  front,  tapering  toward  the  head ;  a  short 
distance  back  of  the  head  expanding  into  a  number  of  irregular,  transparent, 
dilated  folds,  which  border  both  sides  of  an  opaque  central  portion,  in  which  two 
longitudinal  canals  are  faintly  outlined.  First  segments  about  three  times  as 
broad  as  long;  median  segments  square,  or  broader  than  long;  ultimate  segments 
nearly  square,  sometimes  broader  than  long,  sometimes  longer  than  broad.  Geni- 
tal apertures  marginal,  opening  a  very  little  in  front  of  the  middle. 

"A  single  specimen  of  this  species  of  Taenia  was  obtained  from  the  intestine 
of  the  Common  Eel  {Anguilla  vulgaris)  August  26,  1885.  The  length  of  the 
specimen,  when  stretched  out  by  fastening  one  end  with  a  needle  to  the  bottom 
of  the  dissecting  dish  and  removing  all  kinks  and  curves  with  a  fine  brush,  was 
170  mm.  The  length  of  the  same  specimen,  after  having  been  preserved  in  alco- 
hol, is  less  than  90  mm.  The  specimen  when  first  obtained  and  placed  in  sea-water 
was  quite  active.  The  body  was  constantly  throwing  itself  into  sinuous  curves, 
while  the  head  and  neck  were  jerked  from  side  to  side  with  a  moderately  rapid 
motion.  In  addition  to  these  movements  the  neck  and  anterior  portions  of  the 
body  constantly  changed  their  shape  by  the  inflation  or  dilatation  of  the  investing 
membranes  into  wide  transparent  folds,  constricted  at  irregular  int-ervals  by  nar- 
row transverse  bands.  The  neck,  meanwhile,  was  alternately  stretched  out  and 
contracted  like  the  body  of  a  Nemertean.  The  anterior  end  of  the  head  pro- 
truded into  a  proboscis  like  papilla.  The  breadth  of  the  head  itself  varied  from 
0.17  to  0.35  mm. 

"In  the  alcoholic  specimen  the  dilatable  folds  of  the  neck  are  much  con- 
tracted and  broken.  They  lie  in  rough,  ragged  frills  along  each  side  of  the  dark 
central  part  of  the  strobile.  The  head  is  truncate  or  blunt  in  front.  The  neck 
immediately  behind  the  sucking-disks  is  almost  as  wide  as  the  head,  flat,  thin, 
and  little,  if  at  all,  tapeving. 

"The    following   measurements    were    made    on    the   living    specimen.      The 


133]  PROTEOCEPHALIDAE—LA  RUE  133 

head  and  neck  changed  their  position  and  shape  so  rapidly  that  it  was  with  the 
greatest  difficulty  that  trustworthy  measurements  could  be  made: 

Millimeters. 

"Breadth  of  head  0.28 

Diameter  of  acetabula  0.12 

Diameter  of  neck,  narrowest  part  0.20 

Distance  of  first  segments  from  head  17.00 

Length  of  fourth  segment  from  end  of  strobile  1.30 

Breadth  of  same,  posterior  end  1.50 

Breadth  of  same,  anterior  end  1.60 

Length  of  posterior  segment  0.90 

Breadth  of  same,  posterior  end  0.60 

Breadth  of  same,  anterior  end  1.25 

"Habitat. — Common  Eel  (Anguilla  vulgaris)  ;  intestine ;  Wood's  Hole,  Mass., 
August  26,  1885;  one  specimen." 

"Von  Linstow  (Compend.  der  Helminth.,  1878)  records  but  two  Taenia  from 
the  Common  Eel,  T.  macrocephala  Creplin  and  T.  hemispherica  Molin.  T.  dilatata 
is  very  different  from  the  former.  Diesing  (Revis.  der  Ceph.,  Ab.  Cycl.,  p.  378) 
mentions  the  latter,  but  gives  no  enumeration  of  characters.  I  do  not  have  access 
to  Molin's  paper,  and  cannot,  therefore  say  whether  T.  dilatata  is  identical  with 
his  species  or  not.  The  peculiar  inflated  character  of  the  neck  suggests  T.  ambigua 
Dujardin,  but  the  difference  in  size  between  the  adult  specimens  is  alone  sufficient 
to  render  the  union  in  the  same  species  impossible." 

Again  Linton  (1897:425)  reported  this  species  from  A.  chrysypa 
thus: 

"Taenia  dilatata  Linton. — No.  4812  U.  S.  N.  M.  From  common  eel  (An- 
guilla chrysypa).  Several  strobiles  but  in  bad  state  of  preservation,  no  scolices; 
anterior  ends  have  been  exceedingly  long  and  slender.  The  characteristic  dilata- 
tions of  this  species  can  not  be  proved  from  these  specimens.  The  general  ap- 
pearance of  the  strobiles,  as  well  as  the  character  of  the  segments,  however,  agrees 
with  this  species. 

"It  may  be  added  that  the  segments  of  these  specimens  agree  with  Molin's 
description  of  his  T.  hemispherica.  With  the  evidence  at  hand,  however,  the 
writer  does  not  feel  justified  in  uniting  the  two  species  of  T.  dilatata  and  T. 
hemispherica." 

Professor  H.  B.  Ward  secured  this  material,  No.  4812,  U.  S.  N.  M. 
for  my  examination  but  its  state  of  preservation  was  too  poor  to  permit 
any  positive  determination  being  made, 

Linton  (1901:435)  recording  the  parasites  of  Anguilla  chrysypa 
reported  this  species  again.  '^Taenia  dilatata  Linton.  Specimens  of 
this  genus  also  taken  in  1899,  three  on  August  2.    Dimensions  in  milli- 


134  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [134 

meters:    Length,  8,  diameter  of  head,  0.28.    Diameter  of  sucker,  0.08. 
Segments  not  mature.    One  specimen  August  28 ;  length  14  mm. ' ' 

It  is  the  writer's  opinion  that  the  folds  of  the  neck  are  of  no  real 
diagnostic  value.  Such  folds  may  be  seen  on  many  other  specimens  of 
cestodes  in  certain  states  of  expansion  and  contraction.  Unfortunately 
Linton  failed  to  figure  or  describe  any  of  those  internal  structures  which 
are  of  real  service  to  the  modern  systematist.  His  drawing  of  the  ma- 
ture proglottid  is  such  that  it  cannot  be  interpreted  but  his  drawings  of 
the  head  and  of  the  last  proglottids  are  of  value  for  comparative  pur- 
poses. Linton  himself  believed  that  the  segments  of  his  species  agreed 
with  Molin's  description  of  T.  hemisphaerica.  Riggenbach  (1896)  con- 
sidered Linton's  T.  dilatata  to  be  identical  specifically  with  T.  hemis- 
phaerica Molin.  Nufer  (1905)  sought  to  show  that  these  two  species 
were  identical  with  Proteocephalus  macrocephalus  (Creplin).  This  may 
prove  to  be  the  case  but  since  the  writer  has  not  been  able  to  study  any 
material  of  the  T.  hemisphaerica  he  has  preferred  to  consider  the  latter 
as  a  separate  species.  He  is,  however,  ready  to  state  that  Linton's  Tae- 
nia dilatata  is  specifically  identical  with  Proteocephalus  macrocephalus 
(Creplin).  T.  dilatata  is  therefore  a  synonym  of  the  last  named  species 
and  it  should  be  dropped  from  the  list  of  valid  names  for  Proteocepha- 
lus species.  Professor  H.  B.  Ward  very  kindly  secured  for  study  Lin- 
ton's slides  and  some  alcoholic  material  of  his  Taenia  dilatata.  These 
specimens,  judging  from  the  statements  in  his  letter  to  Professor  Ward, 
must  be  from  the  lots  described  by  Linton  in  1889  and  in  1901.  Meas- 
urements of  the  heads,  suckers,  proglottids  and  the  cirrus-pouch,  to- 
gether with  the  general  appearance  of  the  worms,  give  ample  reason  for 
considering  this  form  to  be  identical  with  specimens  taken  from  An- 
guilla  chrysypa,  Lake  Sebago,  Maine,  and  which  the  writer  is  describing 
as  P.  macrocephalus.  These  specimens  agree  in  minute  details  of  struc- 
ture with  the  cestodes  which  Schneider  (1905)  described  as  P.  macro- 
cephalus. A  more  extended  discussion  of  this  comparison  will  be  made 
in  the  proper  connection  (Vide  infra). 

Olsson  (1893)  reported  specimens  of  cestodes  from  Anguilla  vulga- 
ris in  Scandinavia.  Riggenbach  (1896)  considered  the  form  to  be  a 
species  of  Ichthyotaenia.  Stossich  (1897:7)  reported  this  species  from 
Anguilla  vulgaris  at  Narenta,  and  again  in  a  later  paper  (1898a  :113-114) 
he  gave  a  short  diagnosis  of  specimens  taken  from  the  same  host  species 
at  Trieste.  In  this  diagnosis  nothing  of  the  inner  anatomy  of  the  worm 
is  given.    His  diagnosis  reads : 

"Lunghezza  220  mm. ;  larghezza  3.5  mm.  Scolice  in  continuazione  del  collo, 
allungato,  ingressato  al'innanzi,  con  grandi  ventose  globose,  situate  anteriormente ; 
rostello  cortissimo,  ottuso,  provveduto  di  una  piccola  ventosa  apicale.    Collo  corto. 


135]  PROTEOCEPHALIDAE—LA  RUE  135 

Proglottidi  prime  corte  e  ottuse,  le  seguenti  subquadrate.  Aperture  genitali  irrego- 
larmente  alterne ;  pene  corto,  sottilissimo,  pendente,  con  una  strozzatura  verso 
I'apice.  Uova  con  due  invogli,  lunghe  0.089  vam.  Rara  nell'  intestino  dell'  An- 
guilla  vulgaris  (Trieste)." 

Stossich  was  evidently  describing  P.  macrocephalus  and  not  the  P. 
hemisphaericus  (Molin).  Miihling  (1898:37,  69)  noted  this  parasite  in 
A.  vulgaris  in  East  Prussia  but  gave  no  description.  Schneider  (1903: 
29)  reported  from  an  eel  a  single  specimen  of  what  he  then  identified 
as  Ichthyotaenia  hemisphaerica.  Later  in  a  footnote  (1905:10)  he  says 
of  these  specimens  "Ichthyotaenia  hemisphaerica  kommt,  wie  es  scheint, 
in  den  Aalen  des  Finnischen  Meerbusens  garnicht  vor.  Das  Exemplar 
welches  ich  friiher  einmal  irrtiimlich  als  /.  hemisphaerica  bestimmt  habe 
(Beitrage  zur  Kenntnis  der  Helminthenfauna  des  Finnischen  Meerbu- 
sens. Acta  Soc.  pro  Fauna  et  Flora  Fennica  26,  No.  3.  1903.  S.  29), 
erwies  sich  bei  genauerer  Unterschung  auch  als  Ichthyotaenia  macro- 
cephala. ' ' 

From  this  statement  the  writer  infers  that  Schneider  at  that  time 
considered  P.  hemisphaericus  to  be  a  distinct  species.  In  the  text  of  the 
same  article  Schneider  (1905:17-19)  described  P.  macrocephalus,  re- 
cording for  the  first  time  in  the  history  of  the  species  a  careful  study 
of  its  inner  anatomy.  For  this  reason  a  condensed  resume  of  his  de- 
scription is  here  given. 

Proteocephalus  macrocephalus:  This  species  is  a  close  relative  of 
P.  percae  and  is  differentiated  therefrom  by  the  very  short  cirrus-pouch 
and  the  short  cirrus.  The  scolex  has  a  diameter  of  about  0.30  mm.  The 
diameter  of  the  large  deep  suckers  is  0.100  mm.  The  fifth  sucker  as  in 
P.  percae  is  a  rudimentary  structure  made  up  of  long  cells  and  having 
a  diameter  of  0.025  mm.  (The  writer  has  shown  that  this  organ  in 
P.  percae  is  a  true  sucker.)  The  neck  is  quite  broad  and  is  strongly 
compressed  dorsoventrally.  The  body  attains  a  length  40  cm.  and  more. 
A  specimen  11  cm.  long  and  1.8  mm.  broad  had  about  200  segments. 
The  youngest  proglottids  are  about  five  times  as  broad  as  long.  The 
middle  mature  proglottids  are  broader  than  long  while  ripe  proglottids 
are  longer  than  broad. 

The  ovaries  appear  from  the  surface  as  two  small  stripes,  as  they 
do  in  P.  percae.  In  old  ripe  proglottids  they  are  of  irregular  shape, 
bent  and  somewhat  overlapped.  The  cirrus  and  vagina  open  irregu- 
larly, alternating  right  and  left,  in  the  middle  or  a  little  anterior  to  the 
middle  of  the  proglottid  margin.  The  cirrus-sheath  scarcely  equals  Yq 
the  proglottid  breadth,  often  much  less.  The  ductus  ejaeulatorius 
(cirrus-canal)  forms  some  convolutions  in  the  proximal  end  of  the 
cirrus-sheath  and  here  widens  out  into  a  vesicula  seminalis,  0.030  mm. 


136  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [136 

in  diameter.  The  convolutions  of  the  vas  deferens  outside  of  the  cirrus- 
sheath  are  excentric  on  the  porose  side.  In  other  species  with  a  long 
cirrus-sheath  this  convoluted  mass  lies  in  the  middle  of  the  proglottid. 
The  vagina  is  provided  with  a  small  sphincter  muscle  close  to  the 
vaginal  opening.  "Without  broadening  out  into  a  reeeptaculum  seminis 
the  vagina  passes  into  the  ovarial  region.  The  oocapt  is  very  muscular. 
The  uterine  passage  measures  0.02  mm.  in  diameter  and  the  vitelline 
reservoir,  0.30  mm.  The  vitellaria  are  voluminous  and  follicular.  The 
opening  of  the  uterine  passage  and  the  well  preformed  uterine  opening 
on  the  ventral  side  are  found  almost  in  the  same  transverse  section 
posterior  to  the  middle  of  the  proglottid.  The  uterus  has  about  eight 
branches  on  either  side.  Testes,  numbering  about  100  in  each  proglottid, 
lie  irregularly  in  one  or  two  layers.  They  measure  about  0.150  by  0.075 
mm.  Ventral  excretory  vessels  measure  about  0.009  mm.  in  diameter, 
the  dorsal  vessels  about  0.0025  mm.  This  species  occurs  only  in  An- 
guUla  vulgaris  and  only  in  small  numbers. 

In  this  connection  the  writer  thinks  it  of  interest  to  note  that 
Creplin's  specimens  were  taken  at  Greifswald  in  Prussia  while  Schnei- 
der's were  taken  in  Finland.  Geographically  these  habitats  are  not 
distant  nor  do  any  barriers  exist  to  prevent  the  movement  of  the  eels 
from  one  locality  to  the  other.  For  this  reason  and  because  of  anatomi- 
cal similarities  it  is  extremely  probable  that  Schneider's  specimens  were 
of  the  same  species  as  Creplin's.  It  is  to  be  noted  that  Dujardin's 
observations  on  the  size  of  specimens  taken  from  Anguilla  vulgaris  at 
Rennes,  France,  agree  very  well  with  those  which  Schneider  describes. 
Nufer  (1905)  described  and  figured  what  he  considered  to  be  P.  macro- 
cephala  (Creplin).  He  found  two  specimens  in  the  intestinal  tract  of 
a  single  Anguilla  vulgaris  from  Lake  Lucerne.  Nufer  shows  such  strik- 
ing inconsistencies  in  his  statements  and  these  agree  so  poorly  with  the 
findings  of  Schneider  and  the  writer  that  it  seems  best  to  analyse  parts 
of  his  paper.  Compare  his  description  of  the  head  (Nufer  1905:137) 
and  his  statement  aboul  the  head  in  his  summary  (Nufer  1905:145) 
with  the  facts  presented  in  tabular  form  on  pp.  146-147  of  his  paper 
where  he  compares  P.  macrocephalus,  P.  ocellatus,  P.  longicollis,  and  P. 
torulosus.    All  of  these  species  he  reports  from  Lake  Lucerne. 

He  writes  (page  137)  : 

"Der  Scolex  ist  sehr  gross,  zietnlich  breit,  rundlich,  und  besitzt  einen  Durch- 
messer  von  0.1512  mm.  Gegen  den  Hals  ist  er  scharf  und  deutlich  abgesetzt  imd 
iibertrifft  alle  ubrigen  im  Vienvaldsidttersee  vorkommenden  Proteocephalensco- 
lices  an  Grosse  urn  Bedeutendes  (The  italics  are  mine,  La  Rue.),  weshalb  Creplin, 
auch  auf  den  Gedanken  gekommen  sein  mag,  diesen  Proteocephalen  mit  "inacro- 
cephaluy'  zu  bezeichnen.     Vorne  ist  der  Skolex  mit  vier  grossen,  runden  Saug- 


137]  PROTEOCEPHALIDAE—LA  RUE  137 

napfen  versehen,  die  ihm  die  wuchtige  Gestalt  verliehen,  wie  ich  sie  sonst  bei 
keinem  Proteocephalen  mehr  angetrofFen  habe.  Je  nach  der  Kontraktion  der 
Saugnapfe  erscheint  ihr  Rand  kreisrund  oder  unregelmassig  gelappt.  Der  aussere 
Durchmesser  derselben  betragt  0.069  mrn,  der  innere  Durchmesser  0.0414  mm. 
Da  der  Skolex  bei  meinem  Praparate  etwas  geschrumpft  ist,  war  es  mir  unmoglich, 
einen  fiinften,  scheitelstandigen  Saugnapf  nachzuweisen,  wie  ihn  von  Linstow 
beobachtet  haben  will;  jedoch  ist  sicher,  dass  eine  Scheitelvertiefung  vorhanden 
ist." 

Also  in  his  conclusions  note  this  (p.  145)  :  "Der  Skolex  iibertrifft  die 
iibrigen  Proteocephalenskolices  an  Grosse;  dementsprechend  ist  aucb 
der  Durchmesser  der  Saugnapfe  grosser." 

In  the  tabulated  data  (pp.  146-147)  Nufer  gives  the  dimensions  of 
the  scolices  and  of  the  suckers  of  the  four  species  thus:  breadth  of 
scolex  of  P.  macrocephalus  0.151  mm.,  P.  ocellatus  0.114  mm.,  P.  lon- 
gicolUs  0.43  mm.,  P.  torulosus  0,255  mm.;  diameter  of  suckers  of  P. 
macrocephalus  0.069  mm.,  P.  ocellatus  0.05  mm.,  P.  longicollis  0.12-0.19 
mm.,  P.  torulosus  0.1  mm.  According  to  these  figures  the  diameter  of 
the  suckers  and  the  breadth  of  the  head  of  P.  macrocephalus  exceed  the 
same  dimensions  as  given  for  P.  ocellatus  only.  It  is  smaller  than  either 
of  the  other  two  with  which  he  compares  it. 

Again  note  in  the  last  sentence  of  the  first  paragraph  quoted  Nufer 
states  that  he  could  not  find  a  fifth  sucker,  yet  in  his  tabulated  data 
(p.  146)  he  gives  the  diameter  of  the  fifth  sucker  as  0.026  mm.  In  this 
he  must  be  quoting  from  von  Linstow 's  description.  Von  Linstow  was 
the  first  to  mention  a  fifth  sucker  for  this  species.  Evidently  Nufer 's 
statements  must  lack  accuracy.  Nufer 's  drawings  are  of  little  help  in 
giving  a  clear  idea  of  the  systematic  position  of  his  specimens. 

Consider  now  his  description  of  the  male  reproductive  organs  (pp. 
140-142)  and  carefully  compare  his  description  with  the  writer's  figure 
(Fig.  57). 

"Der  Cirrusbeutel  ist  keulenformig;  vorne  an  der  Mundung  schmal,  erweitert 
er  sich  nach  hinten  ziemlich  rasch  und  erhalt  einen  grossten  Durchmesser  von 
0.094s  mm.  Am  Hinterende  ist  er  abgeru^^det  und  geht  bis  uber  die  Mitte  des 
Gliedes  hinaus.  Wie  von  Linstow  (1875)  am  Cirrus,  so  beobachtete  ich  am 
aussern  Drittel  des  Cirrusbeutels  eine  Einschnurung,  durch  die  er  ungefahr  die 
von  V.  Linstow  angegebene  Gestalt  eines  Kegels  erhalt  wie  sie  beim  Kegelspiel 
ublich  ist.  Er  dringt  nicht  vollkommen  gerade  in  das  Innere  des  Gliedes  ein, 
sondern  richtet  sich  etwas  schrag  gegen  das  Hinterende  der  Proglottids.  Die 
Wandung  des  Cirrusbeutels  wird  von  einer  kraftig  ausgebildeten  Muskulatur 
umgeben.  Aehnlich  wie  bei  P'roteocephalus  ocellatus  heften  sich  an  die  Basis  des 
Cirrusbeutels  kraftig  entwickelte  Retraktoren,  welche  die  biischelige  VVurzelfasern 
an  einem  Bulbus  sitzen.  Der  Raum  zwischen  der  Wandung  des  Cirrusbeutels 
und    Cirrus    ist    von    zarten    Bindegewebszellen    und    zahlreichen,    unregelmassig 


138  .ILLINOIS  BIOLOGICAL  MONOGRAPHS  [138 

\^rteilten  Parenchymkemen  erfullt;  vielleicht  sind  auch  einzelne  kleine  Kalk- 
korperchen  eingestreut,  wie  sie  von  Kraemer  (1892)  bei  Proteocephalus  ocellatus 
gefunden  worden  sind. 

"Der  Cirrus  bildet  die  direkte  Fortsetzung  des  Vas  deferens;  er  durchbricht 
den  Cirrusbeutel  am  Hinterende  und  durchzieht  ihn  in  geradlinigem  Verlaufe. 
Nachdem  er  den  hintem  Drittel  des  Beutels  passiert  hat,  nimmt  dessen  Durch- 
messer  ziemlich  rasch  ab  und  reduziert  sich  auf  die  Halfte.  Nach  den  Beobach- 
tungen  Kraemer's  (1892)  ist  der  Cirrus  bei  Proteocephalus  ocellatus  und  P. 
torulosus  in  seinem  hinteren  Teile  in  mehrere  Schlingen  gelegt,  wahrend  er  bei 
P.  macrocephatus  als  vollstandig  gestrecktes  Organ  auftritt.  Gleichbedeutend 
wie  diese  Schlingen  betrachte  ich  einige  ringformige  wulstige  Verdickungen  am 
hintem  Drittel  des  Cirrus.  Der  Schmarotzer  besitzt  in  ihnen  jedenfalk  eine 
Einrichtung,  welche,  ahnlich  wie  der  in  Schlingen  gelegte  Cirrus,  dazu  dient,  bei 
der  Begattung  den  Cirrus  zu  verlangern.  Bei  dem  Geschlechtakte  wiirden  dann 
die  ringformigen  Wiilste  sich  glatten  und  successive  verschwinden  in  dem  Masse, 
als  der  Cirrus  vorgestossen  wird.  Er  ist  von  einer  strukturlosen,  glatten  Wandung 
umgeben,  der  nicht  wie  bei  P.  ocellatus  und  P.  torulosus  nach  hinten  gekriimmte 
Chitinhakchen  aufsitzen.  Sein  Lumen  betragt  im  hintem  Drittel  0.009  mm.,  nimmt 
aber  nach  vorn  bedeutend  ab. 

"Das  Vas  deferens  liegt  hinter  und  vor  dem  Cirrusbeutel  etwas  gegen  den 
Seitenrand  des  Gliedes  verlagert  und  bildet  ein  dichten  Knauel,  der  sich  besonders 
in  der  Lange  der  Proglottis  ausdehnt ;  die  Schlingen,  die  mit  Samenfaden  erftillt 
sind,  greifen  kreuz  und  quer  durcheinander.  Die  Wandung  des  Vas  deferens  ist 
viel  zarter  als  diejenige  des  Cirrus  und  lasst  keine  besondere  Muskulatur  erkennen, 

"Die  Hoden  liegen  als  grosse,  rundliche  Gebilde  im  Mittelfeld  des  Gliedes 
zwischen  den  Dotterstocken,  dem  Keimstocke  und  dem  vordern  Gliedrande. 
Genauere  Angaben  iiber  deren  Zahl  zu  geben,  ist  wegen  vorgeriickter  Reife  der 
Glieder  unmoglich ;  dagegen  durften  sie  ihres  grossen  Durchmessers  (0.054  rnm.) 
wegen  kaum  in  grosserer  Zahl  auftreten.  Bei  Proteocephalus  ocellatus,  dessen 
Hodenblaschen  beinahe  gleichen  Durchmesser  (0.057  nim.)  besitzen,  fand  Kraemer 
(1892)  27-30  in  jeder  Proglottis." 

It  will  be  noted  that  in  nearly  every  particular  his  description 
agrees  with  the  writer's  drawing  (Fig.  57)  and  with  his  description  of 
the  male  organs  of  P.  fallax  La  Rue,  parasitic  in  Coregonus  fera.  The 
work  of  the  writer  on  P.  fallax  was  done  on  material  from  Zschokke-s 
collection,  and  it  seems  that  there  can  be  no  doubt  that  these  specimens 
belong  to  the  species  described  by  Zschokke  and  Kramer  as  Taenia 
ocellata  and  by  Nufer  called  P.  ocellatus  in  his  table.  In  the  length 
and  shape  of  the  cirrus-pouch,  in  the  position  of  the  constriction  of  the 
pouch  and  in  the  inner  bulbous  end  of  the  same  Nufer 's  description  and 
the  writer's  draAving  (Fig.  57)  agree  well.  The  circular  swollen  thick- 
ening which  he  notes  in  the  posterior  third  of  the  cirrus  can  be  best 
interpreted  to  be  a  coil  in  the  ductus  ejaculatorius.  When  such  a  coil 
is  seen  under  certain  conditions  the  appearance  described  by  Nufer  is 


139]  PROTEOCEPHALIDAE—LA  RUE  139 

simulated  almost  exactly.  Likewise  in  the  position  of  the  mass  of  coils 
of  the  vas  deferens,  in  the  number,  size  and  arrangement  of  the  testes 
Nufer's  description  and  his  tabulated  data  (p.  146)  agree  very  well 
with  the  writer's  drawing  and  description  of  the  same  organs  in  P.  fal- 
lax  La  Rue.  His  description  of  the  male  organs  of  the  species  in  ques- 
tion cannot  be  said  to  agree  with  the  writer's  drawing  (Fig.  49)  wliich 
is  a  delineation  of  the  proglottid  of  P.  macrocephalus.  Nor  does  his 
description  agree  with  Schneider's  or  my  own  description  of  the  pro- 
glottid of  that  species. 

Unfortunately  the  writer  has  not  had  the  opportunity  of  examining 
Nufer's  material  of  this  species  nor  any  other  specimens  of  cestodes 
taken  from  the  eels  of  Lake  Lucerne.  He  cannot  therefore  make  a 
positive  determination  of  Nufer's  form.  The  evidence  at  hand  permits 
several  interpretations.  These  are:  1,  that  Nufer  permitted  his  cestode 
material  from  Anguilla  to  become  confused  with  specimens  of  P.  fallax 
from  Coregonus  fera;  2,  that  the  Anguilla  of  Lake  Lucerne  harbor  P. 
fallax;  3,  that  the  ripe  proglottids  described  by  Nufer  had  been  taken 
into  the  intestine  of  the  eel  with  its  food,  perhaps  Coregonus  fera,  liv- 
ing or  dead.  Supposition  1  is  always  a  possibility  when  one  is 
working  with  such  material.  Supposition  2  does  not  give  complete 
satisfaction  because  if  the  eels  of  Lake  Lucerne  harbor  P.  fallax  Nufer 
ought  to  have  been  able  to  find  a  fifth  sucker,  which  he  failed  to  do. 
The  third  supposition  permits  one  to  consider  that  Nufer's  whole  speci- 
mens were  really  P.  macrocephalus  but  that  the  ripe  proglottids  (in 
reality  from  P.  fallax)  upon  which  he  based  his  description  of  the  pro- 
glottids became  confused  with  the  specimens  of  P.  macrocephalus  by 
natural  means.  The  embryo  of  Nufer's  form  is  about  the  size  of  the 
embryo  of  P.  macrocephalus  as  determined  by  the  writer.  His  meas- 
urement of  the  embryo  was  0.0184  mm.  while  the  embryos  of  the  writer's 
specimens  of  P.  macrocephalus  measured  0.017-0.019  mm.  and  rarely  as 
much  as  0.021  mm.  in  an  elongated  embryo.  The  embryo  of  P.  fallax 
is  much  larger,  according  to  the  writer's  measurements  being  0.0312- 
0.0336  mm.  in  diameter. 

Thus  far  attention  has  not  been  called  to  Nufer's  description  of 
the  female  generative  organs.  He  claims  that  the  histological  structure 
of  the  vaginae  of  P.  macrocephalus  and  of  P.  ocellatus  differs  in  some 
respects.  The  writer's  own  investigation  of  the  two  species,  especially 
when  the  investigations  of  Kramer  are  taken  into  consideration,  lead 
him  to  believe  that  the  vaginae  of  the  two  species  are  much  alike  in 
histological  detail.  Nufer  describes  the  ovary  as  being  single,  not  bi- 
lobed.  In  this  he  has  been  led  astray  by  a  condition  sometimes  found 
in  ripe  end-proglottids.     In  such  cases  the  lobes  of  the  ovary  may  be 


140  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [140 

pressed  backward  until  the  ends  of  the  lobes  touch  in  the  median  line. 
In  the  investigations  of  the  writer  covering  a  large  number  of  Proteoce- 
phalidae  he  has  been  unable  to  find  a  fusion  of  the  ovarian  lobes.  He 
must  conclude  that  Nufer's  drawing  of  this  condition  is  hardly  trust- 
worthy. Nufer  also  claims  that  the  waUs  of  the  uterine  pouches  event- 
ually degenerate  so  that  the  eggs  come  to  lie  in  the  parenchjTna.  There 
is  reason  to  doubt  the  accuracy  of  this  statement.  Many  sections  of  a 
large  number  of  old  ripe  proglottids  of  P.  macrocephalus  and  other 
species  of  this  genus  have  been  examined  by  the  writer  who  has  failed 
to  find  any  such  degeneration  of  the  uterine  walls.  Nufer's  material 
must  have  been  in  extremely  poor  condition,  or  else  he  wofully  misin- 
terpreted the  appearance  of  the  sections. 

The  foregoing  discussion  of  Nufer's  article  leads  to  the  conclusion 
that  the  complete  specimens  taken  by  Nufer  from  AnguUla  vulgaris 
were  probably  P.  macrocephalus,  as  he  identified  them.  One  must  fur- 
ther conclude  that  a  part  of  his  description  was  based  on  the  two  loose 
end-proglottids  which  really  belonged  to  P.  faUax  and  that  these  were 
ingested  with  the  food  of  the  eel  or  that  they  were  accidentally  put 
with  the  cestodes  of  AnguUla  during  Nufer's  manipulation  of  the  sp>eci- 
mens.  It  also  seems  that  the  eggs  which  Nufer  measured  came  from 
P.  macrocephalus.  Liihe  (1909)  gave  a  very  short  diagnosis  of  this 
form  but  included  no  figures  of  it.  La  Rue  (1911:475)  included  this 
form  in  a  list  of  Proteocephalus  species  and  stated  that  Taenia  dUatata 
was  a  synonym  of  P.  macrocephalus. 

Some  specimens  which  have  been  identified  by  the  writer  as  P. 
macrocephalus  were  collected  by  Professor  H.  B.  Ward  in  the  course  of 
a  biological  investigation  of  -Sebago  Lake,  Maine  during  the  summer  of 
1907.  This  investigation  was  conducted  under  the  auspices  of  the 
United  States  Fish  Commission.  Professor  Ward  examined  11  speci- 
mens of  AnguUla  chrysypa  Baf.  Of  this  number  five  had  no  cestode 
infection  whatever;  two  yielded  two  Proteocephalids  each;  one  yielded 
one  Proteocephalid,  and  two  others  together  yielded  eighteen  cestodes 
and  some  pieces.  Of  these  eighteen  cestodes  eight  were  Proteocephalids. 
Some  of  the  pieces  also  belonged  to  this  group.  Altogether  then  there 
were  thirteen  Proteocephalids  plus  some  pieces  but  there  was  no  com- 
plete strobUa  among  them.  The  specimens  described  are  from  bottles 
No.  47,  54,  56,  71  and  72  of  the  Sebago  Lake  collection. 

The  worms  are  long  and  slender.  No  complete  strobila  was  found 
but  a  fragment  from  the  middle  region  of  a  worm  measured  120  mm. 
long.  The  maximum  breadth  observed  was  1.8  mm.  Young  proglottids 
are  much  broader  than  long,  measuring  0.33  mm.  broad  by  0.022  mm. 
long.     Mature  proglottids  are  broader  than  long,  measuring  1.0-1.20 


141]  .  PROTEOCEPHALIDAE—LA  RUE  141 

mm.  broad  by  0.40-0.63  mm.  long.  The  length  of  the  ripe  proglottids 
may  exceed  the  breadth.  They  measure  0.90-2.08  mm.  long  by  0.491- 
1.20  mm.  broad.  The  head  (Fig.  1)  is  very  short.  It  has  a  slightly 
rounded  apex  which  is  not  marked  by  grooves  or  furrows.  The  head  is 
somewhat  flattened  dorsoventrally,  being  0.320  mm.  broad  at  the  base 
of  the  suckers  and  about  0.150-0.160  mm.  thick.  At  or  slightly  above 
its  broadest  zone  the  head  bears  the  four  almost  spherical  suckers  which 
have  a  maximum  diameter  of  0.095-0.106  mm.  The  opening  of  the 
sucker  measures  0.050-0.060  mm.  Tho  not  measured  the  sucker  cavity 
is  fairly  deep.  The  musculature  of  the  suckers  is  moderately  well 
developed. 

A  fifth  sucker  is  not  present  in  this  material,  nor  did  Schneider  or 
Nufer  find  one.  It  is  replaced  by  a  structure  such  as  was  described  by 
the  writer  for  P.  filaroides  and  which  was  there  called  an  endorgan. 
This  structure  is  in  reality  a  vestigial  sucker.  It  has  no  elongated  cells, 
tho  Schneider  says  that  he  saw  such  cells,  and  there  is  no  trace  of 
muscles.  In  my  specimens  it  is  composed  of  a  very  few  cells  of  irregu- 
lar shape  surrounded  by  a  membrane  that  is  homologous  with  a  basal 
membrane  in  other  suckers.  This  vestigial  fifth  sucker  is  situated  a  few 
micra  below  the  cuticula  and  it  has  no  connection  with  the  exterior.  It 
measures  0.020-0.025  mm.  This  structure  does  not  resemble  the  fifth 
sucker  of  P.  percae  for  it  has  been  shown  (Figs.  8,  9,  120)  that  in  P. 
percae  this  is  a  muscular  organ  which  doubtless  functions  as  a  sucker. 

The  neck  is  long  and  broad  and  thin.  Frequently  it  measures 
0.225-0.300  mm.  or  more  in  breadth  while  its  length  varies  from  5.0  to 
7.0  mm.  The  strobilation  is  quite  evident.  The  notches  between  the 
segments  are  deep  and  the  angles  of  the  proglottids  are  somewhat 
rounded.  Many  shallow  longitudinal  folds  give  the  surface  of  the  worm 
a  roughened  appearance.  The  genital  aperture  is  marginal,  situated 
about  the  middle  of  the  proglottid,  and  is  irregularly  alternating.  There 
is  no  genital  papilla.  The  excretory  and  nervous  systems  have  not  been 
carefully  investigated.  The  musculature  of  the  strobila  is  much  as 
Benedict  (1900)  found  it  for  P.  amhloplitis  tho  the  muscles  are  not  as 
heavily  developed. 

The  testes  (Fig.  49)  lie  in  a  single  layer  in  the  dorsal  region  of  the 
medullary  parenchyma  between  the  vitellaria  and  anterior  to  the  ovary. 
They  are  closely  packed  together  until  many  of  them  are  more  or  less 
polygonal  in  shape.  They  measure  0.063-0.074  mm.  long  by  0.042-0.063 
mm.  broad  and  they  number  from  100  to  120.  The  vas  deferens  forms 
a  thick  mass  of  coils  extending  from  the  middle  of  the  proglottid  to  the 
very  short  cirrus-pouch.  Schneider  very  well  says  that  the  mass  of 
coils  of  the  vas  deferens  is  excentric  lying  on  the  side  in  which  the 


142  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [142 

cirrus-pouch  is  situated.  The  cirrus-pouch  (Figs.  47,  48,  49)  is  very- 
short,  reaching  just  a  little  way  through  the  vitellaria.  Its  length  is 
about  0.160  mm.  making  the  ratio  of  its  length  to  the  proglottid  breadth 
about  1 :6  to  1 :8.  The  cirrus-pouch  is  bent  up  toward  the  inner  dorsal 
wall  of  the  dermo-muscular  sac.  Within  the  cirrus-pouch  the  ductus 
ejaculatorius  forms  but  one  or  two  coils  before  passing  over  into  the 
short,  but  thick-walled  cirrus.  The  cirrus  is  straight,  ft  was  not  pro- 
truded in  any  specimens  examined  by  the  writer. 

The  vagina  lies  anterior  to  the  cirrus-pouch,  but  the  opening  of 
the  vagina  into  the  genital  atrium  is  situated  almost  dorsal  to  that  of 
the  cirrus  (Fig.  48).  Very  near  the  opening  of  the  vagina  there  is  a 
weak  sphincter  vaginae.  The  lumen  of  the  vagina  in  its  initial  region 
is  quite  lai^.  This  however  soon  constricts  and  the  remainder  of  the 
vagina  is  of  small  diameter.  The  drawings  (Figs.  47,  48)  show  how  the 
vagina  which  opens  anterior  and  dorsal  to  the  cirrus-pouch  passes  over 
the  latter  in  its  course  to  the  ventral  surface  and  then  backward  to  the 
ovary.  Several  species  of  Proteocephalus,  e.  g.,  P.  percae,  P.  pinguis, 
and  P.  exiguus  show  this  twisting  of  the  vagina  part  way  round  the 
cirrus-pouch.  This  point  has  not  been  investigated  in  many  species  but 
it  seems  probable  that  it  is  quite  a  constant  character  of  the  genus. 

The  broadened  region  of  the  vagina  bears  a  ciliated  lining  on  its 
inner  surface  but  the  presence  of  the  cilia  could  not  be  demonstrated 
throughout  the  length  of  the  vagina.  A  small  receptaculum  seminis 
was  found  a  little  way  anterior  to  the  mid-piece  of  the  ovary.  The 
ovary  (Fig.  49)  is  a  heavj'  bilobed  structure  of  much  the  same  shape  as 
in  P.  percae  or  P.  cernuae  tho  shorter  in  those  species.  The  viteUaria 
are  long  follicular  ma^es  iii  the  lateral  fields-  of  the  proglottids.  The 
follicles  are  closely  compacted.  An  oocapt  possessing  a  heavy  muscula- 
ture is  present,  also  an  oot}i)e  with  its  surrounding  shellgland. 

The  uterus  (Fig.  47)  in  ripe  proglottids  is  made  up  of  a  median 
tube  with  7-11-12-14  lateral  outpocketings  on  either  side.  The  septa 
between  the  uterine  pouches  are  sometimes  very  thin  and  in  other  cases 
quite  thick.  The  uterine  passage  is  a  small  tube  which  after  a  sinuous 
course  empties  into  the  uterus  about  the  middle  of  the  proglottid.  The 
uterine  openings  are  1-2  in  number.  Schneider  in  his  specimens  found 
only  a  single  uterine  pore.  In  reality  this  difference  between  the  find- 
ings of  Schneider  and  the  writer  is  slight  and  too  inadequate  to  serve 
as  a  basis  for  a  specific  distinction-  A  second  uterine  pore  is  very  read- 
ily overlooked.  Especially  is  this  true  in  some  species  in  which  the 
second  pore  when  present  is  smaller  than  the  first  and  is  situated  some 
distance  from  it.  The  eggs  were  taken  from  the  uterus  of  alcoholic 
specimens.    The  thin  outer  hyaline  shell,  characteristic  of  the  e^s  of 


1431  PROTEOCEPHALIDAE—LA  RUE  143 

many  species,  could  not  be  demonstrated.  The  outer  shell,  in  this  case 
corresponding  to  the  second  shell  of  most  Proteocephalus  species,  was 
thick  and  quite  granular.  It  measured  0.026-0.036  mm.  in  diameter, 
0.026-0.029  mm.  being  the  more  common  dimensions.  The  embryo,  itself, 
varies  from  spherical  to  ovoidal.  The  diameter  varies  from  0.017  to 
0.019  mm.  and  rarely  0.021  mm.  for  a  long  oval  embryo. 

The  identity  of  this  species  with  Linton's  T.  dUatata  has  already 
been  discussed.  Whether  it  is  the  same  as  P.  hemisphaericus  cannot 
now  be  determined.  A  discussion  of  its  relationship  to  that  form  is 
given  under  the  description  of  P.  hemisphaericus.  Proteocephalus  mac- 
rocephalus  shows  no  marked  resemblances  to  any  other  of  the  North 
American  species  of  Proteocephalus.  In  some  respects  it  shows  some 
marked  resemblances  to  P.  cernuae  La  Rue  but  it  differs  from  that 
species  in  lacking  a  fifth  sucker,  in  the  possession  of  many  more  and 
larger  testes,  in  having  a  cirrus-pouch  that  is  much  shorter  actually  and 
relatively,  in  having  a  much  more  voluminous  mass  of  vas  deferens  and 
in  having  smaller  embryos.  The  proportions  of  the  proglottids  are  also 
different.  While  in  some  respects  this  species  resembles  P.  percae  its 
lack  of  a  fifth  sucker  readily  differentiates  these  two  species.  Moreover 
its  very  short  cirrus-pouch  is  in  marked  contrast  with  the  very  long 
cirrus-pouch  of  P.  percae.  This  species  is  very  different  from  P.  torulo- 
siis  in  size,  in  the  arrangement  of  the  testes,  in  the  size  of  the  cirrus- 
pouch  and  in  the  number  of  uterine  pouches. 

PROTEOCEPHALUS  AMBLOPLITIS  (Leidy) 
[Figs.  18,  19,  116,  117,  134a  &  b,  183] 

1887 :  Taenia  amhlopUtis 

1887:  Taenia  micropteri 

1896 :  Ichthyotaenia  amhlopUtis 

1897:  Taenia  ocellata 

1900 :  Proteocephalus  amhlopUtis 

1905 :  Proteocephalus  amhlopitis 

1909 :    Proteocephalus  sp. 

1911 :    Proteocephalus  amhlopUtis 

Specific  Diagnosis:  Characters  of  the  genus.  Cestodes  large,  280- 
410  mm.  long,  2.-2.5  mm.  in  maximum  breadth.  Surface  of  body  rough, 
with  transverse  and  longitudinal  furrows.  Scolex  prominent,  0.57-0.88 
mm.  broad,  divided  into  quadrants  by  deep  grooves.  Tip  of  scolex 
usually  marked  by  a  minute  depression  with  a  slight  elevation  in  the 


Leidy 

1887 :22-23 

Leidy 

1887 :23 

Riggenbach 

1896 :267-268 

Linton 

1897 :425-426 

Benedict 

1900:339:355 

Marshall  and 

Gilbert 

1905 :513-522 

La  Rue 

1909:21,25, 

27,  28,  31, 36 

La  Rue 

1911:475 

144 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


[144 


center.  Suckers  large,  round  or  oval  in  outline,  deep,  directed  outward 
and  anteriad.  Length  of  suckers  0.300-0.400  mm.  Fifth  sucker  vesti- 
gial, deeply  embedded  in  tissue  of  head.  Proglottids  closely  joined  to- 
gether, corners  rounded.  Young  proglottids  12-15  times  broader  than 
long.  Mature  and  ripe  proglottids  broader  than  long,  about  quadrate, 
or  rarely  longer  than  broad. 

Genital  sinus  irregularly  alternating,  marginal,  situated  about  end 
of  first  fourth  of  the  proglottid.  Vagina  opening  anterior  to  cirrus- 
pouch.  Vaginal  sphincter  long  and  very  thick.  Receptaculum  seminis 
present.  Uterus  with  15-20  lateral  outpocketings  on  either  side.  Cirrus- 
pouch  pyriform,  muscular,  0.400-0.500-0.650  mm.  long,  reaching  ^/y-Vs 
across  the  proglottid  breadth.  Cirrus,  when  protruded,  long  and  slen- 
der. Ductus  ejaculatorius  much  coiled.  Vas  deferens  forming  a  great 
mass  of  coils  extending  to,  or  past,  the  middle  of  the  proglottid.  Testes 
70-100  in  number,  situate  between  vitellaria.  Eggs  provided  with  three 
membranes.  Outermost  membrane  hyaline,  ellipsoidal,  0.036-0.043  mm. 
in  diameter;  second  membrane  thick,  granular,  0.022-0.024  mm.  in 
diameter;  third  closely  investing  the  embryo;  embryo  0.0168-0.018  mm. 
in  diameter. 

Habitat :    Intestine  of  host. 


Host 

Locality 

Collector 

Authority 

Ambloplites  rupestris 

Lake  George,  N.  Y. 

Jos.   Leidy 

Leidy 

(type  host) 

(type  locality) 

1887 :22-23 

Micropterus  Salmoides  = 

Lake  George,  N.  Y. 

Jos.   Leidy 

Leidy 

nigricans 

(type  locality) 

1887 -^s 

Ambloplites  rupestris 

J.  W.  Milner 

Linton 

1897  ■■425-426 

Micropterus  dolomieu 

Lake  St.  Clair,  Mich. 

H.  B.  Ward 

Benedict 
1900 :339-355 

Micropterus  dolomieu 

Lake  Mendota,  Wis. 

Marshall  & 

Marshall  & 

Gilbert 

Gilbert 
1905  ■.513-522 

Micropterus  salmoides 

Walnut  Lake,  Mich. 

T.  L.  Hankinson 

La  Rue 

(The  present  paper) 

Micropterus  salmoides 

Pelican  Lake,  Minn. 

E.  G.  Davis 

La  Rue 

(The  present  paper) 

Amia  calva 

Lake  Erie 

H.  B.  Ward 

La  Rue 

(The  present  paper) 

Atnia  calva 

Lake  St.  Clair,  Mich. 

H.  B.  Ward 

La  Rue 

(The  present  paper) 

145]  PROTEOCEPHALIDAE—LA  RUE  145 

This  species  was  first  described  by  Leidy  (1887:22-28).  His  speci- 
mens were  secured  from  the  stomach  of  Amhloplites  rupestris,  Lake 
George,  New  York.    His  diagnosis  reads: 

"Taenia  AmblopHtis. — Head  quadrate,  spheroidal,  consisting  almost  entirely 
of  the  four  large  spherical  bothria,  with  the  summit  slightly  prominent  and  conical 
or  depressed  and  unarmed ;  neck  very  short  or  none ;  body  compressed  cylindrical, 
gradually  widening  from  the  head  to  near  the  posterior  part,  where  it  slightly 
narrows  to  the  end ;  segments  linear,  becoming  gradually  longer  and  wider,  and 
the  more  quadrate,  all  deeply  and  pretty  regularly  wrinkled  into  two  or  three 
annuli.  Genital  apertures  obscure.  Length  8  to  12  inches;  in  alcohol  contracted 
to  2%  to  5  inches ;  greatest  width  2  mm. 

"Head  0.5  to  0.625  mm.  long  and  0.75  to  0.875  broad.  Bothria  0.375  mm. 
diameter.  Commencement  of  body  0.625  wide.  Anterior  segments  0.125  long,  0.625 
wide;  subsequently  0.375  long  and  1.5  to  1.875  wide;  posterior  segments  0.75  long 
by  I  mm.  wide. 

"A  number  of  specimens  from  the  stomach  of  the  Rock  Bass  Ambloplites 
rupestris.    Lake  George,  New  York. 

"This  species  resembles  the  Taenia  ocellata  Rudolphi  of  the  European  Perch, 
Perca  fluviaiilis,  and  perhaps  is  the  same." 

Leidy 's  drawings  of  the  head  and  proglottids  are  reproduced  (Figs. 
134a  and  b).  Immediately  following  his  report  on  Taenia  amhloplitis 
Leidy  (1887:23)  reported  Taenia  micropteri  from  the  intestine  of  Mi- 
cropterus  salmoides  (nigricans)  Lake  George,  N.  Y.  m  the  following 
words : 

"Taenia  Micropteri. — Head  large,  compressed  spheroidal,  with  four  subter- 
minal  spherical  bothria  and  a  papilliform  unarmed  summit ;  neck  none ;  body  ob- 
scurely segmented,  and  with  no  obvious  internal  organs,  posteriorly  variably  nar- 
rowed and  obtusely  rounded  at  the  end.  Length  from  half  an  inch  to  an  inch, 
and  about  i  mm.  wide.  Apparently  a  larval  form ;  found  in  the  body  cavity  of 
the  Black  Bass,  Micropterus  nigricans.  Six  worms,  soft,  white,  and  active.  The 
longer  ones  of  an  inch  would  elongate  to  double  the  length,  becoming  proportion- 
ately narrower.  The  head,  about  i  mm.  or  more  in  diameter,  varied  in  length  and 
breadth,  according  to  contraction,  sometimes  one  and  sometimes  the  other  being 
the  larger.     Lake  George,  N.  Y." 

This  form  is  probably  the  larval  form  of  P.  amhloplitis.  In  dimen- 
sions and  in  general  character  the  two  forms  are  very  much  alike,  even 
to  the  papilliform  unarmed  summit  of  the  head.  The  obscure  segmen- 
tation was  probably  due  to  wrinkling,  not  to  a  true  segmentation.  P. 
anibloplitis  is  the  only  adult  Proteocephalid  reported  from  Micropterus 
dolomieu  and  M.  salmoides  and  it  is  known  to  occur  in  a  closely  related 
host  species  in  the  same  lake  in  which  Leidy  found  Taenia  micropteri. 
This  is  additional  evidence  in  support  of  the  view  that  Leidy 's  species 


146  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [146 

are  identical.  In  the  light  of  the  evidence  the  name  Taenia  micropteri 
is  to  be  considered  a  synonym  of  Taenia  amhloplitis  and  it  should  be 
stricken  from  the  list  of  valid  species  because  of  the  priority  of  the 
latter  named  species. 

Eiggenbach  (1896:267-268)  put  this  species  in  his  list  of  Ichthyo- 
taenia  yet  he  considered  it  a  doubtful  form.  Under  the  name  Taenia 
ocellata  Rud.  Linton  (1897)  described  this  species  from  Ambloplites 
rupestris.  His  data  agree  very  well  with  the  data  of  Leidy,  Benedict, 
and  the  writer.  A  difference  in  the  length  of  the  neck  caused  Linton 
to  doubt  the  identity  of  his  form  with  Leidy 's  T.  amhloplitis.  This 
difference  is  easily  explained  in  forms  so  contractile  and  so  wrinkled  as 
these.  Linton  himself  says,  "My  specimens  agree  with  Doctor  Leidy 's 
pretty  well  except  in  the  character  of  the  neck.  In  T.  amhloplitis  the 
neck  is  described  as  'short  or  none'.  In  my  specimens  the  neck  is  long." 
Again  in  concluding  Linton  writes,  ''This  reference  of  these  Taenia  of 
Amhloplites  to  the  species  T.  ocellata  is  provisional  only.  I  think,  how- 
ever, that  there  can  be  little  doubt  but  that  my  specimens  are  identical 
with  T.  amhloplitis  Leidy.  The  apparent  absence  of  neck  in  Leidy 's 
species  may  be  ascribed  to  the  presence  of  strong  transverse  wrinkles, 
due  probably  to  the  action  of  the  preserving  fluid."  Benedict  (1900: 
339-355)  working  in  Dr.  H.  B.  Ward's  laboratory  redescribed  this  spe- 
cies using  Leidy 's  specimens  for  comparison  with  material  which  Pro- 
fessor Ward  had  collected  from  Micropterns  dolomieu  while  engaged  in 
a  biological  investigation  of  Lake  St.  Clair  in  August,  1893.  Benedict 
by  means  of  the  section  method  demonstrated  that,  despite  some  dis- 
crepancies in  size,  his  specimens  and  Leidy 's  agreed  in  anatomical  and 
histological  details.  Marshall  &  Gilbert  (1905:  513-522)  found  this  spe- 
cies in  Micropterns  dolomieu  but  not  in  other  hosts  from  Lake  Mendota, 
Wis. 

La  Rue  (1909:21  et  seq.)  referred  to  a  cestode  found  in  Microp- 
teris  salmoides  as  P.  sp.  and  in  a  footnote  on  the  same  page  (p.  21) 
says,  "This  form  may  prove  to  be  Proteocephalus  amhloplitis  Leidy, 
which  it  very  much  resembles".  That  species  is  now  known  to  be  P. 
amhloplitis  Leidy.  These  specimens  are  referred  to  in  the  present  arti- 
cle as  No.  TLH947  and  No.  TLH1036,  collected  by  Hankinson.  La  Rue 
(1911:475)  included  this  species  in  a  list  of  species  of  Proteocephalus. 

The  following  description  of  this  species  is  based  partly  on  Bene- 
dict's (1900)  description  and  largely  upon  the  writer's  own  work  on 
material  which  Professor  T.  L.  Hankinson  had  sent  to  Professor  Ward 
for  determination.  This  material  was  collected  from  the  intestine  of 
Micropterns  salmoides  in  the  course  of  a  biological  investigation  of 
Walnut  Lake,  Michigan,  summer  of  1906.     Other  specimens  have  been 


1471  PROTEOCEPHALIDAE—LA  RUE  147 

examined  and  identified  as  belonging  to  this  species  as  follows:  in  un- 
numbered bottle,  collection  of  La  Rue,  labelled  "Black  Bass,  Large 
mouth.  Pyloric  Region,  Pelican  Lake  Minn.,  1909,"  collected  by  E.  6. 
Davis;  in  bottle  No.  10.151  labelled  "Ex  dogfish,  Sandusky  0.  Jy.  7, 
1905";  in  bottle  No.  2d  labelled  "7/10/01.  P.  in  B.  Host.  Amia  calva- 
Intestine" ;  in  bottle  No.  85  "7/12/01.  P.  in  B.  Amia  calva.  Intestine" ;  in 
bottle  No.  295,  "8/4/01.  P.  in  B.  Amia,  Stomach."  The  abbreviation 
*P.  in  B.'  stands  for  Put-in-Bay,  Ohio,  the  locality  of  collection.  The 
bottles  numbering  10.152,  2d,  85,  295  are  in  Professor  Ward's  collection. 
These  together  with  the  material  collected  by  Hankinson  were  turned 
over  to  the  writer  for  study  through  the  kindness  of  Professor  Ward. 

This  is  one  of  the  largest  of  the  species  of  Proteocephalus  and  it 
is  the  largest  species  of  that  genus  known  at  present  from  the  fishes  of 
North  America.  It  may  attain  a  length  of  41  cm.  and  maximum  breadth 
of  2.0-2.5  mm.  The  body  is  thick,  fleshy  and  heavily  muscled.  In  pre- 
served specimens  the  outlines  of  the  worm  are  somewhat  rough,  due  to 
the  furrows  on  the  margins  of  the  strobila.  The  surface  of  the  worm  is 
also  rough.  Longitudinal  and  transverse  furrows  are  frequent.  Espe- 
cially noticeable  is  a  deep  median  ventral  furrow.  On  account  of  these 
furrows  the  strobilation  is  more  or  less  indistinct.  Likewise  the  poste- 
rior limit  of  the  neck  is  frequently  obscured  by  numerous  transverse 
folds  which  give  an  appearance  of  segmentation. 

The  head  is  large  and  prominent.  Its  breadth  is  0.75-0.875  mm. 
according  to  Leidy,  0.82-0.88  according  to  Benedict.  The  heads  exam- 
ined by  me  measured  0.57-0.60  mm!  Benedict  describes  the  shape  of  the 
head  thus: 

"From  the  anterior  face  it  presents  a  nearly  square  outline,  with  a  deep 
notch  in  the  middle  of  each  side,  dividing  the  surface  into  quarters.  Each  quarter 
contains  a  large  sucker  which  is  directed  outward  and  upward.  The  apex  of  the 
head  is  a  smooth,  rounded  prominence  with  a  small  depression  in  the  top.  No 
hooks  are  present.  A  fairly  good  idea  of  the  shape  of  the  scolex  can  be  obtained 
by  placing  two  truncated  pyramids  base  to  base.  One  of  the  smaller  bases  will 
represent  the  beginning  of  the  neck,  the  other  the  prominence,  while  on  the  slope 
just  beneath  this  would  lie  the  suckers.  The  notches  seen  in  the  anterior  view  of 
the  scolex  form  furrows  down  the  four  sides  of  the  head,  which  gradually  de- 
crease in  depth  and  vanish  on  the  first  few  proglottids." 

Frequently  the  head  is  more  globose  than  Benedict  describes  it.  Such 
heads  are  figured  (Figs.  18,  19).  The  small  apical  papilla  may  not 
always  be  seen  but  the  furrows  have  been  present  in  all  heads  examined 
by  me.  Leidy 's  figure  of  the  head,  reproduced  (Fig.  134a),  is  somewhat 
similar  to  mine.  It,  however,  does  not  show  the  furrows.  Two  of  Bene- 
dict's figures  of  the  head  have  been  reproduced  (Figs.  116,  117)  for 
purposes  of  comparison. 


148  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [148 

The  suckers  which  are  deep  and  well  muscled  measure  0.30  mm.  in 
length  with  a  sucker  opening  of  0.150  mm.  in  my  specimens.  Leidy's 
measurement  of  the  suckers  was  0.375  mm.  long  and  Benedict's  was 
0.32  mm.  broad  by  0.40  mm.  long.  In  suckers  of  this  size  differences  in 
contraction  states  will  explain  the  variation  in  size.  There  is  no  func- 
tional fifth  sucker  but  deep  in  the  tissue  of  the  head  a  peculiar  structure 
(Fig.  116)  may  be  found  which  is  a  vestigial  fifth  sucker.  Benedict 
saw  the  structure  but  was  unable  to  point  out  its  significance.  The  neck 
is  short  and  0.25-0.45  mm.  wide.  The  first  proglottids  are  much  broader 
than  long  measuring  0.344  mm.  broad  by  0.022-0.028  mm.  long.  Mature 
and  ripe  proglottids  are  broader  than  long  as  a  rule,  but  at  times  may 
be  quadrate,  or  even  longer  than  broad.  The  last  condition  is  rare. 
The  maximum  breadth  of  mature  and  ripe  proglottids  is  1.5-2.0-2.5  mm. 
while  the  maximum  length  of  the  same  is  0.5-1.0-2.0  mm. 

The  cuticula,  the  musculature,  and  the  nervous  system  were  care- 
fully investigated  by  Benedict  (1900)  who  found  nothing  especially  re- 
markable about  these  structures.  The  excretory  system  differs  from 
that  of  P.  fossatus  (Rigg.)  and  Corollabothrium  lohosum  Rigg.  in  hav- 
ing no  definitely  placed  foramina  secundaria.  At  irregular  interv^als 
along  either  dorsal  or  ventral  excretory  vessel  arise  numerous  branches 
which  take  a  fairly  direct  course  to  the  exterior.  These  branches  are 
of  various  sizes  and  the  lumen  of  each  undergoes  considerable  variation 
in  size  in  its  course.  The  duct  through  the  cuticula  is  very  small  and 
not  beset  with  small  bristles  as  Kraemer  (1892)  described  for  the  species 
which  he  determined  to  be  P.  filicollis.  No  special  musculature  can  be 
found  in  the  course  of  these  ducts  to  the  exterior.  The  main  longi- 
tudinal excretory  vessels  are  four  in  number.  They  are  situated  just 
within  the  dermo-muscular  sac.  The  ventral  vessel  has  the  larger  lumen 
and  it  is  further  differentiated  from  the  dorsal  vessel  by  its  verj'  thin 
membranous  lining.  The  dorsal  vessel  has  a  lining  apparently  made  up 
of  columnar  epithelial  cells.  Rarely,  however,  the  dorsal  vessel  is  di- 
lated to  a  size  equalling  that  of  the  ventral  vessel  and  in  these  cases  the 
structure  of  the  walls  of  the  two  vessels  appears  to  be  identical.  No 
transverse  vessel  connecting  the  ventral  or  dorsal  vessels  in  the  poste- 
rior part  of  the  proglottid  has  been  found.  In  the  scolex  there  is  an 
intricate  plexus  of  excretory  vessels.  Many  ducts  opening  to  the  exte- 
rior by  small  pores  place  the  coils  of  this  plexus  in  communication  with 
the  exterior. 

The  common  marginal  genital  sinus  opens  about  one  fourth  of  the 
length  of  the  proglottid  from  the  anterior  end.  It  is  irregularly  alternate 
in  position.  A  genital  papilla  is  not  present.  The  vagina  opens  into  the 
sinus  anterior  to  the  cirrus-pouch.    The  testes  are  spheroidal  bodies, 


149]  PROTEOCEPHALIDAE—LA  RUE  149 

0.050-0.065  mm.  in  diameter,  which  are  sometimes  rendered  polyhedral 
by  pressure  of  adjacent  testes.  They  number  from  75  to  100  and  are 
crowded  into  one  or  two  irregular  layers  in  the  space  between  the 
vitellaria  and  anterior  to  the  ovaries.  The  vas  deferens  (Fig.  183) 
forms  a  large  mass  of  coils  beginning  in  the  mid-field  of  the  proglottid 
and  reaching  to  the  cirrus-pouch.  A  large  part  of  this  mass  of  vas 
deferens  is  posterior  to  the  cirrus-pouch.  The  cirrus-pouch  (Fig.  183) 
is  a  large  pyriform  structure,  heavily  muscled  with  both  longitudinal 
and  circular  muscles.  Its  larger  and  inner  end  is  attached  by  heavy 
muscle  strands  to  the  dermo-museular  sac.  It  measures  0.40-0.50-0.65 
mm.  long  by  0,150-0.230  mm.  in  maximum  breadth.  The  ratio  of  its 
length  to  the  proglottid  breadth  varies  from  2:7  to  2:5.  When  it  is 
protruded  the  cirrus  is  long  and  slender,  a  little  larger  at  the  base  than 
at  the  tip  but  when  within  the  cirrus-pouch  the  basal  part  of  the  cirrus 
has  a  broad  lumen.  This  sharply  defines  the  cirrus  from  the  slender 
ductus  ejaculatorius  which  forms  an  intricate  mass  of  coils. 

The  vagina  (Fig.  183)  always  lies  anterior  to  the  cirrus-pouch. 
Very  near  its  opening  the  vagina  dilates  markedly  and  in  this  region 
it  is  provided  with  a  thick  and  powerful  sphincter  vaginae  measuring 
about  0.030  mm.  thick  by  0.300  mm.  long.  This  prominent  sphincter 
surrounding  the  greatly  dilated  vagina  renders  the  latter  a  striking 
feature  of  the  proglottid  and  makes  a  valuable  diagnostic  character  for 
the  species.  At  the  inner  limits  of  the  sphincter  the  vagina  contracts 
sharply  and  beyond  the  constriction  it  again  dilates.  Here  its  inner 
surface  is  weakly  ciliated.  Anterior  to  the  ovary  the  vagina  may 
describe  a  few  coils  before  it  passes  into  that  portion  of  its  length  which 
is  differentiated  into  a  small  receptaculum  seminis.  Entering  the  inter- 
ovarial  space  the  vagina  describes  a  few  coils  and  then  discharges  into 
the  oviduct.  The  ovary  (Fig.  184)  is  situated  in  the  posterior  part  of 
the  proglottid.  It  is  bilobed,  each  lobe  being  thick,  broad  and  somewhat 
club-shaped.  The  lobes  extend  laterad  to  the  vitellaria.  From  the  mid- 
piece  of  the  ovary  arises  a  muscular  organ,  the  oocapt  or  gulping  organ. 
From  the  oocapt  arises  the  oviduct  which  after  making  one  or  more 
coils  is  joined  by  the  vagina.  From  this  point  the  oviduct  extends  to 
the  ootype.  Just  as  it  enters  the  ootype  the  oviduct  receives  the  common 
vitelline  duct  which  has  been  formed  by  the  union  of  the  paired  vitelline 
ducts  arising  in  the  posterior  part  of  the  vitellaria. 

The  ootype  is  surrounded  by  the  so-called  shell-glands,  the  indi- 
vidual cells  of  which  are  long  and  clubshaped.  The  ootype  discharges 
into  a  somewhat  muscular  duct  known  as  the  uterine  passage  which 
extends  anteriad  passing  the  ovary  on  the  dorsal  side.  It  discharges  into 
the  uterus  from  the  dorsal  side  of  the  latter  at  a  point  about  0.080-0.110 


ISO  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [150 

mm.  anterior  to  the  ovary.  Benedict  (1900)  stated  that  there  was 
no  uterine  passage  in  this  species.  An  examination,  however,  of  his 
slides  and  reconstruction  drawings  convinced  the  w^riter  that  a  uterine 
passage  was  present  in  Benedict's  specimens  as  it  was  in  his  own.  The 
vitellaria  are  lateral  and  follicular.  In  this  species  they  do  not  extend 
posterior  to  the  lobes  of  the  ovary  nor  do  they  parallel  the  posterior 
margin  of  the  proglottid.  The  uterus  in  young  proglottids  is  a  median 
ventral  tube  from  which  later  15-20  lateral  outpocketings  develop  on 
either  side.  Benedict's  drawing  reproduced  (Fig.  183)  shows  but  a 
part  of  the  pouches.  There  are  1-2-3  preformed  uterine  pores.  The 
uterine  eggs  are  covered  with  three  membranes,  an  outer,  thin  hyaline,  a 
middle  thick  and  granular  membrane  and  an  inner  thin  membrane 
closelj  investing  the  embryo.  The  outer  membrane  which  is  not  spher- 
oidal but  ellipsoidal  measures  about  0.036-0.043  mm.  in  length.  The 
middle  shell  is  spherical  0.022-0.024  mm.  in  diameter.  The  embryo 
measures  0,0168-0.017  mm.  in  diameter. 

The  American  forms  which  most  resemble  this  species  are  P.  per- 
plexus  LaRue  and  P.  nematosoma  (Leidy).  The  latter  may  be 
the  same  as  P.  amhloplitis.  P.  perplexus,  however,  is  distinguished 
from  P.  ambloplitis  by  its  smaller  head,  smaller  suckers,  thin- 
ner and  smaller  strobila,  by  the  smaller  size  and  the  different  loca- 
tion of  its  sphincter  vaginae,  by  the  posterior  prolongation  of  its 
vitellaria,  by  its  much  smaller  cirrus-pouch,  by  its  fewer  coils  of  ductus 
ejaculatorius,  by  its  larger  number  of  testes,  and  by  the  smaller  size  of 
the  eggs.  P.  ambloplitis  greatly  resembles  P.  nematosoma.  It  differs 
from  that  species  chiefly  in  the  somewhat  larger  size  of  the  head  and 
suckers.  P.  amhloplitis  is  larger  than  most  of  the  old  world  species  of 
Proteocephalus  and  it  differs  from  a  large  number  of  these  in  not  pos- 
sessing a  fifth  sucker.  It  resembles  P.  torulosus  in  the  lack  of  a  function- 
al fifth  sucker  and  in  size.  It  may  be  easily  differentiated  from  P.  toru- 
losus by  reason  of  its  larger  head  and  suckers,  by  its  sphincter  vaginae 
which  for  size  and  length  is  unique  in  the  genus,  by  its  more  numerous 
uterine  pouches,  and  by  the  different  arrangement  of  testes. 

P.  amhloplitis  may  be  distinguished  from  all  other  known  species 
of  Proteocephalus  by  means  of  its  extremely  large  sphincter  vaginae 
which  because  of  its  length  and  its  extraordinary  development  is  re- 
markable. This  species  is  also  readily  distinguished  from  all  other 
species  of  the  genus  by  reason  of  the  large  number  of  coils  of  the  ductus 
ejaculatorius. 


151]  PROTEOCEPHALIDAE—LA  RUE  151 


PROTEOCEPHALUS  PERPLEXUS  La  Rue 

[Figs.  17,  64,  65] 

?  1886 :     Taenia  fiUcollis  Leidy,     1886 :62-63. 

1911 :    Proteocephahis  perplexus  La  Rue,  1911 :478-479. 

Specific  Diagnosis:  The  characters  of  the  genus.  Observed  length 
as  much  as  15.5  cm.  Maximum  breadth  1.7  mm.  Strobilation  evident. 
Angles  of  proglottids  sharp  and  distinct.  First  proglottids  much  broader 
than  long.  Mature  proglottids  broader  than  long,  1.70  mm.  broad  by 
0.595  mm.  long.  Ripe  proglottids  quadrate  or  longer  than  broad,  1.02 
mm.  broad  by  1.10  mm.  long  to  1.75  mm.  long  by  0.510  mm.  broad. 
Head  somewhat  spheroidal,  flattened  dorsoventrally,  divided  into  four 
quadrants  by  grooves  extending  nearly  to  the  apex.  At  apex  frequently 
a  small  papilla  situated  in  a  small  depression.  Head  0.663-0.714  mm. 
broad,  0.425-0.510  mm.  long.  Suckers  four,  0.340-0.459  mm.  long  by 
0.255-0.272  mm.  broad.  Cavity  of  sucker  deep.  No  fifth  sucker,  no 
rostellum.    Neck  broad  and  thick,  about  0.5  mm.  long. 

Genital  pore  marginal,  situated  at  end  of  first  fourth  or  half  of 
proglottid,  irregularly  alternating.  No  genital  papilla.  Cirrus-pouch 
elongated  oval  in  shape,  0.30-0.344  mm.  long,  extending  %-i/4-%  across 
the  proglottid.  Ductus  ejaculatorius  with  1-3  coils.  Cirrus  when  pro- 
truded spindle-shaped,  slender,  thicker  at  base,  0.60  mm.  long.  Testes 
135-155  in  number,  in  one  layer  occupying  dorsal  field  between  vitellaria 
anterior  to  ovary.  Testes  0.069  by  0.037  mm.  in  diameter.  Vagina 
anterior  to  cirrus-pouch,  never  crossing  same,  beginning  region  dilated. 
Inner  surface  of  vagina  heavily  ciliated.  Vitellaria  follicular,  volumi- 
nous, follicles  of  same  large,  compacted.  Portion  of  vitellaria  parallel- 
ling posterior  margin  of  proglottid.  Ovarian  lobes  thick,  heavy,  irreg- 
ular. Uterus,  when  developed,  possessing  20-25  lateral  pouches.  Ute- 
rine pores  2-4.  Uterine  eggs  provided  with  three  membranes.  Inner 
and  outer  membranes  thin,  second  thick,  granular,  sometimes  partially 
or  wholly  split  into  two  layers.  Embryos  0.013-0.014  mm.  by  0.014- 
0.0156  mm.,  second  membrane  0.019-0.030  mm.,  outer  membrane  0.024- 
0.036  mm.  in  diameter. 

Habitat :    In  intestine  of  host. 


152 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


[152 


Host 

Locality 

Collector 

Authority 

Amia  calva  L. 

Illinois   river, 

H.  B.  Ward 

La  Rue  (Present  paper) 

(type  host) 

Havana,  III. 
(type  locality) 

?  Amia  calva  L. 

North  Carolina 

Jos.  Leidy 

La  Rue  (Present  paper) 

Lepisosteus 

Illinois  river 

H.  B.  Ward 

La  Rue  (Present  paper) 

platostomus 

Type:  Alcoholics  No.  Ha89a  and  slides  from  the  same  lot.  Co- 
types  :— Ha  10  a ;  Ha  10  e ;  Ha  29  a ;  Ha  30  h ;  Ha  77  a,  b,  e ;  Ha  94 
e,  h,  and  slides  of  Ha  94  h,  in  Dr.  H.  B.  Ward's  Collection. 

Leidy  (1886:62-63)  found  some  specimens  of  a  cestode  in  Amia 
calva  from  North  Carolina.  These  specimens,  he  thought,  might  be 
Taenia  filicollis  Kud.  His  specimens,  judging  from  his  data  probably 
belong  to  the  species  P.  perplexus  but  since  Leidy  published  no  draw- 
ings of  his  specimens  no  positive  determination  can  be  made.  That 
part  of  his  report  containing  his  data  is  here  quoted : 

"The  worms  accorded  with  the  description  of  the  Taenia  filicollis,  infesting 
Sticklebacks,  Gasterosteus,  and  is  probably  the  same  species.  They  range  from 
iH  to  3  inches  long,  gradually  widening  from  the  delicate  thread-like  neck  to  the 
posterior  rounded  extremity,  where  they  measure  from  i  to  1.5  mm.  wide.  The 
head  is  spheroidal,  variably  broader  or  longer,  and  about  0.625  mm.,  with  the 
summit  slightly  prominent  and  unarmed  and  with  four  hemispherical,  lateral  both- 
ria  0.25  mm.  in  diameter.  Neck  variable,  when  extended  long  and  narrow  and 
usually  about  half  the  width  of  the  head.  Anterior  segments,  transversely  linear, 
about  an  eighth  the  length  of  the  breadth,  gradually  becoming  inverted  saucer- 
shaped  or  scutellate,  and  about  one-fourth  the  length  of  the  breadth.  Posterior 
segments  more  quadrate,  slightly  widening  behind,  about  0.75  mm.  long  and  from 
1  to  1.5  mm.  broad;  last  segment  longest  and  rounded.  Genital  apertures  mar- 
ginal." 

La  Eue  (1911:478-479)  briefly  described  this  species  and  gave  it 
the  name  Proteocephalus  perplexus.  The  specimens  upon  which  this 
species  is  based  were  collected  at  Havana,  111.,  June  and  July  1910,  by 
Dr.  H.  B.  "Ward.  The  hosts  Amia  calva  and  Lepisosteus  platostomus 
were  caught  in  the  Illinois  river  at  that  place.  Four  Amia  were  exam- 
ined, three  were  infested  with  30,10  and  15-20  specimens  of  P.  perplexus 
respectively.  One  was  uninfested.  Eight  gars  were  examined;  three 
were  infested  with  6,  3  and  5  specimens  of  P.  perplexus  respectively. 
Five  of  the  gars  were  not  infested  with  this  species.    In  all  the  Amia 


153]  PROTEOCEPHALIDAE—LA  RUE  153 

and  in  one  of  the  infested  gars  the  P.  perplexus  was  not  accompanied 
by  any  other  species  of  Proteocephalus.  In  two  of  the  gars  P.  singu- 
laris  occurred  together  with  P.  perplexus.  Nothing  is  known  regarding 
the  seasonal  distribution  of  the  parasite  and  almost  nothing  regarding 
its  geographical  distribution.  It  has  not  been  found  in  the  Amia  of  the 
Great  Lakes  which  Professor  Ward  examined  a  number  of  years  ago. 
In  those  specimens  were  found  P.  amhlopUtis. 

Judging  from  external  appearances  alone  these  specimens  of  P. 
perplexus  are  very  much  like  specimens  of  P.  amhlopUtis.  Even  after 
staining  and  mounting  some  pieces  in  toto  they  appear  to  belong  to 
that  species  tho  certain  points  of  difference  may  be  indistinctly  made 
out.  Examination  of  sections  revealed  certain  characters  by  which  the 
two  species  may  be  differentiated.  The  writer  (1911:478)  proposed  the 
name  Proteocephalus  perplexus  for  this  species  by  reason  of  the  diffi- 
culties attending  its  differentiation  from  its  nearest  congener.  In  size 
P.  perplexus  is  somewhat  smaller  than  P.  amhlopUtis.  Its  strobila  is 
shorter,  narrower  and  thinner.  Its  head  is  a  little  smaller  than  that 
of  P.  amhlopUtis  but  is  very  similar  in  shape  and  it  has  four  grooves 
as  does  that  species.  Sometimes  the  minute  papilla  at  the  apex  of  the 
head  thought  to  be  characteristic  of  P.  amhlopUtis  may  be  seen.  The 
suckers  are  large  and  in  a  position  similar  to  that  which  they  occupy  in 
P.  amhlopUtis. 

The  head  (Fig.  17)  is  somewhat  spheroidal.  It  is  flattened  dorso- 
ventrally  and  is  evenly  rounded  anteriorly.  Four  grooves  extend  from 
the  base  of  the  head  nearly  to  the  apex  where  frequently  there  is  a 
small  papilla  in  a  shallow  depression.  The  head  is  not  as  truncate  as 
the  head  of  P.  amhlopUtis  described  by  Benedict  (1900).  It  resembles 
very  closely  heads  which  the  writer  has  examined  and  drawn  (Figs.  18, 
19)  of  the  latter  species.  Four  heads  of  P.  perplexus  measured  in 
breadth  and  length  0.663  by  0.510  mm.,  0.697  by  0.425  mm.,  0.697  by 
0.510  mm.,  0.714  by  0.459  mm.  The  suckers  measured  0.340-0.459  mm. 
long  by  0.255-0.272  mm.  broad.  There  is  no  fifth  sucker  and  no  rostel- 
lum.  Sections  through  the  apex  of  the  head  failed  to  reveal  even  a  vestige 
of  a  fiifth  sucker.  In  P.  amhlopUtis  the  rudimentary  fifth  sucker  is 
large.  The  suckers  are  deep  and  well  muscled,  usually  longer  than 
broad.  The  sucker-opening  is  directed  outward  and  slightly  forward. 
The  neck  is  0.5-0.6  mm.  long.  It  is  broad  and  thick  but  narrower  than 
the  head.  No  complete  strobila  was  found  yet  most  of  the  longer  pieces 
contained  ripe  proglottids.  Six  pieces  with  heads  measured  65,  91,  106, 
105,  125,  and  155  mm.  long  respectively.  The  maximum  breadth  ob- 
served was  1.7  mm. 


154  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [154 

The  worm  shows  a  very  evident  strobilation,  the  angles  of  the  pro- 
glottids  being  sharp  and  well  marked.  The  posterior  border  of  one 
proglottid  overlaps  the  anterior  end  of  the  one  following.  Segmenta- 
tion begins  about  0.5-0.6  mm.  from  the  head.  Here  the  segments  are 
much  broader  than  long.  Gradually  the  length  increases  in  proportion 
to  the  breadth  until  in  nearly  mature  proglottids  the  dimensions  may 
be  1.70  mm.  broad  by  0.595  mm.  long.  Further  down  the  strobila  nearly 
ripe  proglottids  may  be  about  quadrate  1.02  mm.  broad  by  1.10  mm. 
long.  Ripe  proglottids  are  longer  than  broad,  1.75  mm.  long  by  0.510 
mm.  broad.  In  all  but  ripe  proglottids  the  genital  pore  is  not  promi- 
nent but  in  these  long  proglottids  the  pore  is  situated  on  an  eminence. 
Moreover  in  this  part  of  the  strobila  the  posterior  region  of  the  seg- 
ment is  much  broader  than  the  anterior  or  middle  region.  The  neck  is 
0.5-0.6  mm.  long.  It  is  broad  and  thick,  not  as  broad,  however,  as  the 
head. 

The  genital  pore  is  situated  on  the  lateral  margin  at  the  end  of  the 
first  one-fourth  to  one-half  of  the  proglottid.  The  pores  alternate  irreg- 
ularly. In  mature  and  nearly  ripe  proglottids  there  is  no  genital 
papilla  but  in  elongated  ripe  proglottids  a  genital  prominence  is  noted. 
Into  the  common  genital  sinus  both  vagina  and  cirrus  open,  the  vagina 
always  anterior  to  the  cirrus.  The  male  organs  (Fig.  65)  much  resem- 
ble the  male  organs  of  P.  anibloplitis  but  here  again  certain  differences 
exist.  The  cirrus-pouch  in  P.  perplexus  is  not  as  voluminous  nor  as 
long  as  in  P.  amhloplitis.  It  measures  0.300-0.344  mm.  long.  Its  length 
goes  into  the  proglottid  breadth  3-4-5  times  depending  somewhat  upon 
the  state  of  contraction  of  the  proglottid.  "Within  the  cirrus-pouch  are 
1-3  coils  of  ductus  ejaculatorius,  a  much  smaller  number  than  in  P. 
amhloplitis.  The  coils  of  the  vas  deferens  outside  the  cirrus-pouch  are 
very  numerous.  They  form  in  mature  proglottids  a  thick  compact  mass 
extending  from  the  cirrus-pouch  to  the  middle  of  the  segment.  The 
cirrus  itself  is  well  muscled.  When  protruded  it  is  a  long  spindle- 
shaped  organ  slightly  thicker  near  the  base  than  elsewhere.  It  is  nearly 
0.60  mm.  long.  The  testes  are  very  numerous,  135-155  in  number.  They 
are  arranged  in  one  layer,  occupying  the  entire  dorsal  field  anterior  to 
the  ovaries  and  as  far  laterad  as  the  vitellaria,  with  the  exception  of 
the  small  region  taken  up  by  a  portion  of  the  cirrus-pouch  and  the  vas 
deferens.  The  testes  measure  as  much  as  0.069  mm.  long  by  0.037  mm. 
broad,  their  short  axis  Ijang  parallel  to  the  long  axis  of  the  worm. 

The  vagina  (Fig.  64)  in  its  first  part  is  considerably  dilated. 
Throughout  the  full  length  of  this  dilated  region  there  is  a  weak  cir- 
cular musculature.  At  the  end  of  the  dilatation  is  a  strong  sphincter 
vaginae  0,053  mm.  long  and  0.015  mm.  thick.    P.  amhloplitis  has  a  long 


155]  PROTEOCEPHALIDAE—LA  RUE  155 

and  extremely  heavy  sphincter  vaginae  while  in  P.  singularis  the 
sphincter  vaginae  resembles  that  of  P.  perplexus.  Throughout  its 
length  as  far  as  the  receptaculum  seminis  the  inner  surface  of  the  vagina 
is  heavily  ciliated.  The  vagina  does  not  cross  the  cirrus-pouch,  but 
after  reaching  the  mid-field  of  the  segment  it  takes  a  sinuous  course 
posteriad  to  the  interovarial  space.  A  small  receptaculum  seminis  lies 
just  anterior  or  posterior  to  the  mid-piece  of  the  ovary.  The  vagina 
forms  no  coils  anterior  to  the  ovary.  The  vitellaria  (Fig.  64)  are 
voluminous,  the  follicles  are  of  good  size  and  closely  packed  together. 
They  extend  not  only  to  the  extreme  posterior  margin  of  the  proglottid 
but  are  bent  around  and  lie  parallel  to  the  posterior  margin  nearly  to 
the  mid-field.  This  is  particularly  noticeable  in  ripe  proglottids.  The 
paired  vitelline  ducts  pass  through  this  posterior  continuation  of  the 
vitellaria. 

The  ovary  is  bilobed.  The  lobes  are  thick  and  heavy,  with  a  some- 
what irregular  outline  due  to  protuberances.  In  sections  it  is  noted  that 
the  ovary  is  not  a  solid  organ  but  is  made  up  of  smaller  parts  more  or 
less  fused  together.  An  oocapt,  ootype,  shell  gland  and  other  organs 
usually  found  in  the  inter-ovarial  space  in  Proteocephalids  are  found 
here  and  in  the  relations  characteristic  of  the  genus.  The  uterus  (Fig. 
64),  a  median  tube  in  mature  proglottids,  is  made  up  in  ripening  pro- 
glottids of  a  median  tube  and  20-25  narrow  lateral  pouches.  These 
pouches  occupy  the  entire  ventral  field  between  the  vitellaria  and  the 
anterior  and  posterior  proglottid  limits.  There  are  2-4  ventral  uterine 
pores.  The  uterine  eggs  have  three  membranes,  an  inner  one  closely 
investing  the  embryo,  a  middle  membrane,  quite  thick  and  granular 
and  frequently  partially  or  completely  split  into  two  layers,  and  an 
outer  membrane  thin  and  hyaline.  The  embryo  measures  0.013-0.014 
mm.  by  0.014-0.156  mm.,  being  usually  somewhat  elongated  but  at  times 
spheroidal.  The  second  membrane  measures  0.019-0.030  mm.  and  the 
outer  one  0.024-0.036  mm.  in  diameter. 

The  excretory  system  is  made  up  of  four  main  lateral  canals  which 
traverse  the  length  of  the  strobila.  The  two  ventral  vessels  are  larger 
than  the  two  dorsal.  At  intervals  small  branches  arise  from  the  main 
vessels  and  lead  to  the  exterior.  No  transverse  excretory  commissure 
was  observed.  In  the  head  and  neck  the  anastomoses  of  the  excretory 
system  are  very  complex.    The  main  trunks  are  nearly  straight. 

This  species  is  closely  allied  to  P.  amhloplitis  yet  it  differs  from 
that  species  in  size,  in  the  lack  of  a  vestigial  fifth  sucker,  in  the  size  and 
location  of  the  vaginal  sphincter,  in  the  posterior  prolongation  of  the 
vitellaria,  in  the  smaller  cirrus-pouch,  in  the  fewer  coils  of  ductus  ejac- 
ulatorius  within  the  cirrus-pouch,  in  the  greater  number  of  testes  and 


156  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [156 

in  the  size  of  the  eggs.  In  size  P.  perplexus  somewhat  resembles  P. 
singularis.  It  also  resembles  P.  singularis  in  the  shape  and  proportions 
of  its  proglottids  and  in  the  character  of  the  vagina  and  vaginal  sphinc- 
ter. These  two  species  differ  in  the  size  and  shape  of  the  head,  suckers, 
and  neck.  Tho  nearly  of  the  same  size  the  strobila  of  P.  perplexus  is 
the  larger.  P.  perplexus  has  more  numerous  testes,  a  larger  cirrus- 
pouch  and  more  numerous  coils  of  ductus  ejaculatorius.  The  main 
excretory  vessels  of  P.  perplexus  are  straight,  but  they  are  very  sinuous 
in  P.  singularis.  P.  perplexus  differs  from  all  the  European  species  in 
the  position  of  the  sphincter  vaginae  and  in  the  extension  of  the  vitel- 
laria  along  the  posterior  border  of  the  proglottid. 


PROTEOCEPHALUS  SINGULARIS  La  Rue 
[Figs.  24,  25,  39-41,  83-87] 
1911 :    Proteocephalus  singularis  La  Rue,  1911 :479 

Specific  Diagnosis:  Characters  of  the  genus.  Strobila  long  and 
slender.  Observed  length  up  to  170  mm.,  length  probably  as  much  as 
250  mm.  in  complete  individuals.  Maximum  breadth  up  to  0.90-1.0  mm. 
Head  small,  0.250-0.30  mm.  broad  by  0.20-0.22  mm.  long.  Head  bear- 
ing at  broadest  part  four  large  suckers.  Apical  region  of  head  fre- 
quently prolonged  into  an  unarmed  rostellum-like  organ.  Deep  grooves 
between  suckers.  Suckers  of  variable  shape,  always  with  pointed  apex, 
with  shallow  cavity  and  thin  muscular  wall.  Length  of  suckers  0.130- 
0.170  mm.,  breadth  of  same  Q.170-0.190  mm.  No  fifth  sucker,  no  vestige 
of  same.  Neck  slender,  2.0-3.0  mm.  long  by  0.1-0.2  mm.  broad.  First 
proglottids  broader  than  long,  0.255  mm.  broad  by  0.017-0.020  mm. 
long.  Mature  proglottids  as  much  as  0.85  mm.  broad  by  0.34-0.37  mm. 
long.  Ripe  proglottids  longer  than  broad  or  quadrate  0.680  mm.  broad 
by  0.90-1.00  mm.  long.  Old  spent  proglottids  up  to  2.0  mm.  long  by 
0.4  mm.  broad.  Segmentation  evident.  Posterior  angles  of  proglottids 
slightly  projecting. 

No  genital  papilla.  Genital  pore  marginal,  irregularly  alternating, 
situated  at  end  of  first  i/4-%  of  proglottid.  Testes  numerous,  75-80  or 
90,  in  a  single  layer  occupying  entire  field  between  vitellaria  anterior 
to  ovary.  Breadth  of  testes  0.4-0.6  mm.,  length  of  same  0.07-0.10  mm. 
Vas  deferens  a  large  mass  of  coils  in  the  mid-field.  Ductus  ejaculato- 
rius with  few  or  no  coils.  Cirrus  slender,  straight,  muscular.  Cirrus- 
pouch  slender,  nearly  straight,  muscular,  0.185-0.20-0.265  mm.  long. 
Length  of  cirrus-pouch  21^-3  times  into  proglottid  width. 


157]  PROTEOCEPHALIDAE—LA  RUE  157 

Vagina  always  anterior  to  cirrus-pouch,  never  crossing  the  latter. 
Beginning  region  of  vagina  narrow.  This  region  about  0.1  mm.  long, 
terminated  by  weak  sphincter  vaginae.  Adjacent  to  narrow  region  a 
dilated  portion.  Vagina  not  ciliated  in  any  part  of  length.  Vesicula 
seminalis  present.  Vitellaria  with  large  follicles,  not  paralleling  pos- 
terior margin  of  proglottid.  Uterus  when  fully  developed  with  20-25 
lateral  outpocketings  on  either  side.  ^  Uterine  pores  ventral,  2-3-4  in 
number.  Eggs  with  three  membranes.  Embryos  0.014-0.0156-0.0168 
mm.,  second  membrane  0.026-0.031  mm.,  outer  membrane  0.027-0.033 
mm.  in  diameter.  Excretory  system  has  four  main  lateral  trunks. 
Ventral  vessels  large,  dorsal  vessels  much  smaller.  All  main  vessels 
sinuous  or  spiral.    Many  secondary  openings  to  exterior. 

Habitat:  In  intestine  of  Lepisosteus  platostomus  (type  host),  Illi- 
nois river,  Havana,  Illinois  (type  locality). 

Type:  Alcoholics  No.  Ha  30  i  and  slides  from  same.  Cotypes, 
Ha  12  d,  e ;  Ha  29  f ;  Ha  87  a ;  Ha  88  a ;  Ha  101b ;  Ha  110c.  Mate- 
rial in  Dr.  H.  B.  "Ward's  collection. 

La  Rue  (1911:479)  in  a  preliminary  way  described  this  species. 

In  a  collection  of  endoparasites  secured  by  Dr.  H.  B.  Ward  from 
fish  caught  in  the  Illinois  river  at  Havana,  Illinois,  were  some  speci- 
mens of  Proteocephalus  somewhat  resembling  the  species  which  the 
writer  (1911)  has  designated  as  P.  perplexus.  On  account  of  the  pecu- 
liar form  of  the  head  and  suckers  of  this  species  the  name  Proteoceph- 
alus singularis  La  Rue  has  been  proposed  for  it.  In  all,  eight  gars  were 
examined.  Of  this  number  only  one  was  uninfested  with  this  parasite. 
This  gar  was  infested  only  with  P.  perplexus.  The  numbers  of  P.  sin- 
gularis present  in  each  of  the  seven  infected  hosts  were :  2,  7,  26,  2,  5, 
4,  8.  Two  gars  had  a  mixed  infection  of  P.  singularis  and  P.  perplexus. 
Slides  have  been  prepared  from  some  of  this  material. 

Its  study  showed  that  the  longest  piece  measured  170  mm.  Com- 
plete strobilas  perhaps  measure  up  to  250  mm.  or  more.  Its  maximum 
breadth  is  about  0.90-1.00  mm.  The  strobila  is  slightly  smaller  than 
that  of  P.  perplexus  yet  it  greatly  resembles  the  latter  species  in  out- 
ward appearance.  The  more  slender  neck  and  the  smaller  head  are 
its  more  readily  noted  points  of  difference.  The  head  (Figs.  24,  25)  is 
small,  measuring  from  0.250  to  nearly  0.300  mm.  in  breadth  by  0.20- 
0.22  in  length.  At  its  broadest  part  it  bears  four  large  suckers  which 
when  well  expanded  cover  up  nearly  the  whole  surface  of  the  head 
except  the  apical  region.  The  apical  region  in  most  species  is  flattened 
or  conical,  but  in  this  species  it  is  drawn  out  into  a  long  slender  pro- 
tuberance which  has  no  hooks.    This  is  not  a  rostellum.    At  times  this 


158  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [158 

protuberance  is  greatly  contracted.  Between  the  suckers  are  deep 
grooves  which  in  many  cases  cause  the  head  to  appear  almost  lobate. 
The  suckers  themselves  are  more  or  less  variable  in  shape  but  they 
always  show  the  pointed  apex.  The  cavity  is  shallow,  the  muscular 
wall  thin.  The  suckers  measure  0.130-0.170  mm.  long  by  0.170-0.190 
nmi.  broad.  A  fifth  sucker  is  not  present.  A  careful  search  through 
three  series  of  sections  has  failed  to  reveal  any  trace  of  an  endorgan. 
The  neck  is  long  and  slender,  2.0-3.0  mm.  to  the  first  traces  of  segmenta- 
tion. The  breadth  is  0.1-0.2  mm.,  usually  much  narrower  than  the  head. 
The  neck  grades  over  almost  imperceptibly  into  the  first  proglottids 
which  are  very  difficult  to  discern. 

The  first  proglottids  are  12-15  times  broader  than  long,  being  about 
0.255  mm.  broad  by  0.017-0.020  mm.  long.  As  the  proglottids  near 
maturity  they  increase  greatly  in  size  measuring  up  to  0.85  mm.  broad 
by  0.34-0.37  mm.  long.  Proglottids  in  which  the  uterus  contains  some 
eggs  measure  0.51  mm.  long  by  0.867  mm.  broad  or  are  even  quadrate, 
each  dimension  being  about  0.85  mm.  The  ripest  proglottids  become 
longer  than  broad,  measuring  0.680  mm.  broad  by  0.90-1.00  mm.  long 
and  in  rare  cases  of  old  spent  proglottids  the  length  may  be  about  2.0 
mm.  and  the  breadth  about  0.4  mm.  Segmentation  is  evident.  No  lon- 
gitudinal furrows  are  present.  Transverse  furrows  occur  only  at  the 
division  lines  between  the  segments.  The  posterior  angles  of  the  pro- 
glottids project  slightly  beyond  the  anterior  margin  of  the  proglottid 
following. 

This  species  has  not  been  thoroly  investigated  as  to  its  histological 
structure.  However,  certain  points  have  been  worked  out  with  some 
care.  The  cuticula,  the.subcuticula,  the  parenchyma,  the  muscles  of  the 
strobila,  and  the  nervous  system  were  not  seen  to  differ  essentially  from 
similar  structures  in  P.  amhloplitis. 

The  muscles  of  the  head  were  worked  out  in  series  of  transverse 
and  frontal  sections.  The  muscles  of  the  apical  protuberance  of  the 
head  could  not  be  worked  out  with  care  but  that  region  could  be  seen 
to  be  well  supplied  with  muscles  especially  in  the  subcuticular  layer. 
The  peripheral  parts  of  the  apical  region  of  the  head  contain  many 
heavy  muscles  going  up  toward  the  tip.  In  the  middle  and  lower  region 
of  the  suckers  about  0.135  mm.  from  the  tip  of  the  head  heavj-  muscles 
cross  from  one  sucker  wall  to  the  wall  of  the  sucker  opposite  forming 
a  heavy  muscle  cross  (Fig.  40).  Weaker  muscles  cross  the  head  dorso- 
ventrally  and  laterally.  These  muscles  extend  to  the  subcuticular  re- 
gion and  serve  by  their  contraction  to  cause  in  part  the  deep  grooves 
between  the  suckers.  Near  the  upper  limits  of  the  suckers  at  a  depth 
of  about  0.10  mm.  from  the  tip  of  the  head  heavy  muscles  (Fig.  39) 


159]  PROTEOCEPHALIDAE—LA  RUE  159 

connect  the  sucker  wall  with  the  lateral  or  dorso-ventral  surfaces  of  the 
head.  These  probably  serve  a  double  function  of  causing  by  their  con- 
traction the  deep  grooves  between  the  suckers  and  of  pulling  the  adja- 
cent margins  of  the  suckers  together,  thus  broadly  flaring  the  sucker. 
In  this  region  the  muscle-cross  is  weakly  developed.  In  the  lower  part 
of  the  head  (Fig.  41)  just  as  the  suckers  disappear  from  transverse 
sections  the  fibers  of  the  muscle-cross  flare  out  to  secure  a  broad  attach- 
ment on  the  sucker  wall.  In  fact  a  great  many  fibers  dip  down  below 
and  attach  themselves  near  the  lower  outer  margin  of  the  sucker.  At 
the  sides  of  each  sucker  large  groups  of  very  heavy  muscles  may  be 
found  the  attachments  and  courses  of  which  are  best  made  out  in  lon- 
gitudinal sections.  Transverse  fibers,  i.  e.  lateral  and  dorsoventral,  pass 
through  the  muscle-cross  and  assist  in  the  formation  of  a  muscle-star 
(Fig.  41).  Below  this  point  the  muscles  rapidly  assume  the  relations  of 
the  muscles  in  the  strobila. 

The  longitudinal  muscles  of  the  head  are  fairly  well  worked  out. 
Certain  groups  which  were  of  undetermined  character  in  transverse 
sections  can  be  readily  distinguished  in  frontal  sections.  The  large 
groups  of  tangentially  cut  ends  (Fig.  41)  at  either  side  of  the  sucker 
are  in  reality  longitudinal  muscles  which  come  up  from  the  neck  region 
and  find  their  attachment  on  either  side  of  the  sucker  (Fig.  87),  right 
and  left.  Much  weaker  muscle  bands  pass  up  toward  the  tip  of  the 
scolex.  In  the  apical  region  (,Fig.  86)  many  heavy  muscles  attached 
to  the  sucker  wall  pass  out  by  diverging  paths  to  the  subcuticular  area. 
These  probably  control  not  only  a  certain  part  of  the  movement  of  the 
suckers  but  also  the  form  of  the  apex.  The  muscles  of  the  head,  both 
longitudinal  and  transverse,  are  much  more  strongly  developed  than  in 
the  head  of  P.  flaroides,  which  the  writer  has  worked  out  with  great 
care  (La  Rue  1910).  Their  arrangement  in  the  two  forms  is  very 
similar,  tho  certain  groups  of  muscles  present  in  P.  singularis  are  lack- 
ing in  P.  filaroides. 

In  the  head  the  excretory  vessel  may  be  followed  as  coiling  and 
anastomosing  trunks  and  vessels  which  are  much  less  developed  than  in 
P.  amhloplitis  or  P.  perplexus.  In  the  neck  region  the  anastomoses  of 
the  excretory  system  are  much  like  those  in  the  head.  Here  four  main  lat- 
eral vessels  are  to  be  found  with  branches  which  have  a  very  few 
openings  to  the  exterior.  A  little  further  posteriad  the  excretory  ves- 
sels increase  in  size.  This  is  particularly  true  of  the  ventral  vessel. 
Both  ventral  and  dorsal  vessels  Jiave  a  very  sinuous  or  spiral  course 
even  in  the  most  elongated  proglottids.  In  frontal  sections  (Fig.  83) 
the  sinuous  ventral  vessels  appear  as  series  of  oval  or  circular  spaces  in 
the  parenchyma.     In  either  frontal  or  transverse  sections  the  ventral 


160  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [160 

vessel  is  very  prominent  on  aeeoimt  of  its  very  large  lumen.  At  fre- 
quent intervals  branches  may  be  seen  to  be  given  off  from  the  ventral 
vessel  and  a  lesser  number  from  the  dorsal.  Many  of  the  ventral 
branches  communicate  to  the  exterior  by  means  of  small  ducts  through 
the  cuticula.  These  branches  are  more  numerous  and  more  prominent 
than  in  any  other  Proteocephalid  examined  by  the  writer.  One  branch 
opens  quite  regularly  at  the  posterior  angle  of  the  proglottid.  Other 
branches  (by  far  the  largest  number)  open  on  the  ventral  surface.  A 
few  branches  of  the  ventral  vessel  open  on  the  dorsal  surface.  No  duct 
connecting  the  ventral  excretory  ducts  in  the  posterior  part  of  the  pro- 
glottids  has  been  observed,  nor  have  the  relations  of  the  excretory  vessels 
been  observed  in  the  end-proglottid. 

There  is  no  genital  papilla.  The  genital  pore  is  marginal,  irregu- 
larly alternating,  and  situated  at  the  end  of  the  first  third  or  two-fifths 
of  the  proglottid.  Vagina  and  cirrus  open  into  the  common  genital 
sinus,  the  vagina  always  anterior  to  the  cirrus.  In  all  the  general 
features  of  the  sexual  organs  these  are  typical  of  the  genus.  The  testes 
are  ovoidal  or  spheroidal  in  shape,  0.04-0.06  mm.  broad  by  0.07-0.10 
mm.  long.  They  are  arranged  (Fig.  85)  in  a  single  layer  which  fills 
the  dorsal  region  between  the  vitellaria  anterior  to  the  ovaries.  The 
region  in  which  the  coils  of  the  vas  deferens  lie  is  free  from  them.  They 
are  75  to  85-90  in  number.  The  vasa  efferentia  can  very  readily  be 
made  out  just  below  the  layer  of  longitudinal  muscles  of  the  dorsal  side. 
These  empty  into  the  vas  deferens  which  makes  a  large  mass  of  coils  in 
the  middle  of  the  proglottid  and  reaching  over  to  the  cirrus-pouch. 
This  knot  is  relatively  thick  for  it  extends  from  dorsal  to  ventral  wall 
of  the  dermo-muscular  sac*  Entering  the  cirrus-pouch  (Fig.  84)  the 
vas  deferens  which  now  becomes  the  ductus  ejaculatorius  passes  over 
almost  immediately  into  the  straight  cirrus.  The  ductus  ejaculatorius 
is  usually  straight  but  sometimes  it  is  thrown  into  from  one  to  three 
small  coils.  The  cirrus  itself  has  not  been  seen  protruded.  It  is  always 
slender,  straight,  and  muscular.  The  cirrus-pouch  (Fig.  84)  is  slender, 
nearly  straight,  and  quite  muscular.  It  measures  0.185-0.200-0.265  mm. 
long.    Its  length  goes  21/^-3  times  into  the  proglottid  width. 

The  vagina  (Fig.  84)  in  its  first  part  has  a  narrow  lumen.  This 
region  extends  about  0.100-0.110  mm.  from  the  vaginal  opening.  It  is 
characterized  by  its  rich  covering  of  gland  cells  and  its  weak  longitudi- 
nal and  circular  muscles.  The  vaginal  sphincter  which  is  poorly  devel- 
oped is  situated  just  at  the  end  of  this  first  region.  The  sphincter  is 
about  0.025  mm.  long.  Immediately  following  this  region  with  the 
narrow  lumen  is  a  dilated  portion  which  may  extend  clear  to  the  vesic- 
ula  seminalis.     There  is  no  ciliated  region  of  the  vagina.     The  vagina 


161]  PROTEOCEPHALIDAE—LA  RUE  161 

does  not  cross  the  cirrus-pouch  but  lies  anterior  thereto  until  it  comes 
to  the  knot  of  the  vas  deferens  which  it  passes  ventrally.  It  forms  no 
coils  anterior  to  the  ovary  but  may  be  somewhat  sinuous.  The  vitellaria 
(Fig.  85)  are  lateral.  The  follicles  are  large,  and  closely  packed  to- 
gether. In  the  posterior  part  of  the  segment  the  vitellaria  never  extend 
posterior  to  the  ovary  as  in  P.  perplexus.  The  vitelline  ducts  are  as  in 
P.  amhloplitis.  The  lobes  of  the  ovary  are  smaller  than  in  P.  perplexus 
or  P.  amhloplitis.  They  are  more  branched  than  in  either  of  those  spe- 
cies but  this  branched  condition  cannot  be  determined  except  in 
sections. 

The  relations  of  the  oocapt,  ootype,  oviduct,  lower  vagina,  vitelline 
ducts,  and  uterine  passage  are  typical  of  the  genus  and  need  no  expla- 
nation here.  The  uterus  is  a  median  tube  in  mature  proglottids.  In 
ripe  proglottids  (Fig.  83)  the  median  tube  has  20-25  lateral  outpocket- 
ings  on  either  side  which  take  up  the  entire  ventral  field  of  the  segment 
bounded  by  the  anterior  and  posterior  margins  and  by  the  ventral 
excretory  ducts.  Two,  three  or  four  ventral  uterine  pores  have  been 
observed,  tho  in  most  of  the  very  old  proglottids  the  ventral  body  wall 
is  split  from  end  to  end.  The  formation  of  the  ventral  uterine  openings 
is  as  La  Eue  (1909)  described  the  process  in  0.  filaroides.  The  eggs 
very  much  resemble  those  of  P.  perplexus  in  size  but  differ  in  having  a 
thinner  and  more  hyaline  middle  membrane  which  is  never  split  into 
two  layers.  The  embryos  measure  0,014-0.0156-0.0168  mm.  The  second 
membrane  measures  0.026-0.031  mm.  and  the  other  hyaline  membrane 
0.027-0.033  mm. 

This  species  in  the  structure  of  the  head  stands  quite  by  itself. 
Its  proglottids  somewhat  resemble  those  of  P.  perplexus  but  are  smaller. 
In  toto  preparations  the  ventral  excretory  ducts  of  P.  singularis  being 
much  larger  make  a  fairly  easy  means  of  separation.  In  the  number 
of  uterine  pouches  the  two  species  are  much  alike,  but  these  pouches  in 
P.  perplexus  extend  farther  laterad  than  in  P.  singularis.  In  the  posi- 
tion of  the  vaginal  sphincter  the  two  species  are  much  alike  but  the 
beginning  region  of  one  is  dilated,  in  the  other  contracted.  One  has  a 
ciliated  vagina,  the  other  not.  The  cirrus-pouch  of  P.  singularis  is 
more  slender,  the  cirrus  and  ductus  ejaculatorius  are  straighter  and 
more  slender  than  in  P.  perplexus. 


152  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [162 

PROTEOCEPHALUS  SULCATUS  (Klaptocz) 

[Figs.  130,  131,  175,  176] 

1906:    Ichthyotaenia  sulcata  Klaptocz  1906:123-130 

1911:    Proteocephalus  sulcatus       La  Rue  1911:475 

Specific  Diagnosis:  The  characters  of  the  genus.  Cestodes  of 
small  size,  up  to  68  mm.  long.  Maximum  breadth  1.5-2.0  mm.  Scolex 
very  variable  in  form,  unarmed,  without  apical  depression  or  fifth 
sucker,  divided  by  distinct  furrows  into  four  quadrants.  Median  fur- 
rows arise  near  first  proglottids.  Cross-section  of  one  type  of  scolex 
nearly  round,  of  the  other  elliptical.  Breadth  of  scolex  0.46-1.75  mm., 
thickness  of  same  0.67-1.28  mm.  depending  on  contraction  states.  Suck- 
ers, deep  rounded,  one  in  each  quadrant.  Diameter  of  sucker  0.250  mm. 
Neck  none.  First  proglottids  broader  than  long.  Mature  and  ripe  pro- 
glottids quadrate  or  longer  than  broad.  Last  proglottid  rounded  pos- 
teriorly. Maximum  breadth  of  proglottid  1.5  mm.,  maximum  length  of 
same  2.0  mm. 

Grenital  pore  at  end  of  first  third  of  segment.  Cirrus-pouch  about 
0.20-0.25  mm.  long,  extending  about  i/4  across  the  proglottid.  Several 
coils  of  ductus  ejaculatorius  in  cirrus-pouch.  Cirrus  when  protruded 
long  and  slender.  Testes  about  200,  irregularly  arranged  between  vitel- 
laria.  Opening  of  vagina  posterior  to  the  cirrus-pouch.  Lumen  of 
vagina  large.  Ovary  posterior,  bUobed.  Vitellaria  follicular,  lateral. 
Uterus  with  10-12  lateral  out-pocketings  in  ripe  proglottids.  Eggs 
pear-shaped,  circular  in  cross-section.  Diameter  of  same  0.016  mm, 
length  up  to  0.019  mm. 

Habitat:  Polypterus  endlicheri  Heckel  (type  host),  taken  at  Duem 
(tj^pe  locality)  on  the  White  Nile,  and  Clarotes  laticeps  Riippell,  from 
the  White  Nile  in  the  region  of  Khartoum. 

The  material  was  collected  by  Dr.  F.  Werner  in  the  spring  of  1905, 
was  described  by  Klaptocz  (1906),  and  was  listed  by  La  Rue  (1911) 
among  other  species  of  Proteocephalus.  In  the  summary  and  also  in  the 
following  description  the  data  are  taken  from  Klaptocz 's  paper  and 
from  his  drawings. 

A  specimen  from  Clarotes  measured  68  mm.,  one  from  Polypterus 
60  mm.  The  scolex  (Figs.  130,  131)  is  of  very  variable  form,  is  un- 
armed, has  no  apical  depression,  and  it  always  shows  four  very  distinct 
furrows  of  which  the  two  on  the  median  line  are  the  heavier.  These 
divide  the  anterior  face  of  the  scolex  into  four  large  similar  quadrants 
whose  acute  angles  come  together  at  the  tip  of  the  scolex.     The  free 


163]  PROTEOCEPHALIDAE—LA  RUE  163 

margins  of  the  quadrants  often  are  indented  especially  on  the  flattened 
sides.  The  furrows,  especially  the  heavier  median  ones,  arise  in  the 
region  of  the  first  proglottids.  In  each  quadrant  a  deep  rounded  sucker 
is  sunken  which  lies  nearer  the  margin  of  the  scolex  than  to  its  tip. 
Two  types  of  scolex  are  recognized  with  intermediate  forms.  The  one 
is  more  or  less  flattened  on  the  anterior  face.  It  has  a  more  or  less 
round  cross  section.  Its  furrows  are  shallow,  and  its  suckers  are  di- 
rected anteriad  while  the  other  type  has  an  elliptical  transverse  section. 
Its  thickness  is  less  than  the  breadth  of  the  scolex.  The  scolex  itself  is 
much  flattened.  The  furrows  are  deep  and  very  plain,  reaching  nearly 
to  the  tip  of  the  scolex.  The  suckers  are  more  elongated  and  the  sucker 
openings  are  directed  outward.  These  types  of  head  are  to  be  consid- 
ered contraction  states  of  the  same  species  since  intermediate  stages  are 
present.  The  second  type  is  found  in  Polypterus,  and  the  first  type  in 
Clarotes.  The  diameter  of  the  scolex  in  specimens  from  Clarotes  lati- 
ceps  ranges  up  to  1.72  mm,  the  diameter  of  a  sucker  of  the  same  0.250 
mm.  The  diameter  of  the  smallest  scolex  of  this  host  is  0.75  mm.  The 
thickness  of  scolices  from  Polypterus  endlicheri  ranges  from  0.67  to  1.28 
mm.,  the  breadth  from  0.46  to  0.675  mm.  No  neck  is  present.  The  young- 
est proglottids  are  thinner  than  the  head  and  only  slightly  narrower. 
Their  breadth  exceeds  their  length.  Proglottids  increase  in  both  breadth 
and  length  with  age  until  they  are  quadrate  in  form  or  longer  than 
broad.  The  last  proglottid  is  rounded  posteriorly.  The  maximum 
breadth  of  a  proglottid  from  Clarotes  is  1.5  mm,  maximum  length  2.0 
mm.  The  maximum  breadth  of  a  segment  from  Polypterus  is  1.5  mm. 
and  the  maximum  length  of  the  same  1.9  mm. 

Sexual  ripeness  appears  very  early.  The  anlagen  of  genital  organs 
in  stained  specimens  may  be  seen  in  very  young  proglottids  within  two 
scolex  lengths  of  the  head.  Many  proglottids  are  sexually  ripe  before 
becoming  quadrate  in  form.  The  marginal  genital  pore  alternates  irreg- 
ularly. Sometimes  in  as  many  as  ten  to  twelve  or  more  proglottids  the 
genital  sinus  may  be  on  the  same  side  tho  this  condition  is  rarely  true 
in  more  than  five  segments.  The  sinus  (Fig.  176)  is  situated  at  about 
the  end  of  the  first  third  of  the  segment.  The  vagina  opens  posterior  to 
the  cirrus-pouch.  The  cirrus-sheath  (Fig,  176)  containing  the  coiled 
cirrus  (or  cirrus  plus  the  ductus  ejaculatorius)  has  a  pear-shaped  form 
and  lies  perpendicular  to  the  margin  of  the  proglottid.  It  may  extend 
14  across  the  breadth  of  the  proglottid.  The  cirrus-pouch  is  about  0.20- 
0,25  mm,  long  (measured  from  Klaptocz's  figures  which  are  drawn  to 
scale).  The  cirrus  (Fig,  176)  when  protruded  is  very  long  and  slender. 
It  is  thicker  at  the  base  than  at  the  point  and  is  devoid  of  hooks  or 
bristles.    The  testes  are  elliptical,  in  number  about  200.    The  testicular 


164  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [164 

field  is  limited  by  the  -vitellaria,  the  anterior  margin  of  the  segment  and 
the  ovaries.  There  is  no  median  zone  free  from  them  except  where 
they  are  pushed  aside  by  the  developing  uterus.  As  the  uterus  con- 
tinues to  develop  the  testes  are  pushed  well  into  the  lateral  fields. 

The  vagina  (Fig.  176)  which  has  a  large  lumen  runs  to  the  middle 
of  the  proglottid  in  a  more  or  less  sinuous  course,  then  bends  and  passes 
directly  back  to  the  posterior  end  of  the  segment.  In  proglottids  with 
developed  uterus  the  vagina  is  pushed  somewhat  from  the  median  line 
and,  it  seems,  always  toward  the  side  on  which  it  opens.  The  ovary  is 
posterior,  is  bilobed,  and  the  lobes  are  limited  laterally  by  the  vitellaria. 
In  the  end-proglottid  the  lobes  of  the  ovary  are  bent  together.  The 
vitellaria  are  follicular  and  lateral.  On  the  porose  side  the  vitellaria  is 
divided  into  two  parts  connected  by  small  follicles  which  lie  dorsal  and 
ventral  to  the  genital  passages.  The  uterus  (Figs.  175,  176)  is  a  median 
tube  reaching  to  the  anterior  end  of  the  proglottid  from  which  arise 
many  (10-12  in  drawing)  lateral  outpocketings.  These  are  separated 
when  fully  developed  by  thin  tissue-layers.  The  eggs  are  pear-shaped, 
circular  in  cross-section,  0.016  mm.  in  diameter  and  0.029  mm.  long. 

Despite  the  fact  that  Klaptocz  found  an  apparently  intergrading 
series  between  the  types  of  head  mentioned  by  him  it  seems  to  the 
writer  that  there  is  a  possibility  of  his  having  confused  two  forms  of 
very  similar  appearance.  The  writer  in  the  present  work  on  Proteo- 
cephalus  perplexus  and  P.  amhloplitis  found  considerable  difficulty  in 
distinguishing  the  heads.  And  in  certain  respects  the  structure  of  the 
internal  organs  of  the  proglottids  was  remarkably  similar.  So  too  it 
was  found  to  be  difficult  to  distinguish  the  proglottids  of  P.  perplexus 
and  P.  singularis.  This  was  made  especially  difficult  when  both  species 
occurred  in  a  specimen  of  Lepisosieus  platostomus.  The  fact  that  this 
species  seems  to  occur  in  two  such  widely  separated  families  of  fish  is 
a  further  intimation  that  two  species  of  parasite  are  here  included 
under  the  one  name.  Material  from  the  two  hosts  should  be  carefully 
reworked. 


165]  PROTEOCEPHALIDAE—LA  RUE  165 


PROTEOCEPHALUS  PENTASTOMA   (Klaptocz) 
[Figs.  126-129] 

1906:    Ichthyotaenia  pentastoma         Klaptocz        1906:130-133 
1911 :    Proteocephalus  pentastomus     La  Rue  1911 :475 

Specific  Diagnosis:  Characters  of  the  genus.  Cestodes  of  small 
size.  Strobila  up  to  28  mm.  long;  maximum  breadth  of  same  1.19  mm. 
Neck  none.  First  proglottids  many  times  broader  than  long.  Mid- 
proglottids  1.19  mm.  broad,  0.21  mm.  long.  Sixth  from  last  pro- 
glottid 0.59  mm.  broad,  0.21  mm.  long.  Head  large,  oval  in  cross- 
section,  0.975  mm.  broad  by  0.83  mm.  thick.  No  rostellum,  no  hooks. 
Four  large  suckers,  oval  or  round  in  outline,  0.50  mm.  in  diameter, 
situated  in  four  quadrants.  Suckers  separated  from  each  other  by 
longitudinal  furrows.  A  true  fifth  sucker  0.085-0.100  mm.  in  diameter 
situated  at  apex  of  head.  Genital  aperture  about  middle  of  proglottid. 
Vagina  posterior  to  the  cirrus-pouch.  Testes  70-100,  arranged  irregu- 
larly between  vitellaria  with  tendency  toward  formation  of  two  lateral 
fields.  No  data  on  cirrus-pouch  and  vas  deferens.  Vitellaria  lateral,  not 
voluminous.    Ovary  bilobed,  posterior.    Uterus  immature. 

Habitat :    Polypterus  hichir  Geoffr.,  White  Nile  near  Khor  Attar. 

Klaptocz  (1906)  described  this  species  and  it  has  not  since  been 
described  from  the  specimens.  La  Rue  (1911:475)  gave  it  a  place  in  a 
list  of  species  of  Proteocephalus.  A  single  specimen  was  taken  by  Dr.  F. 
"Werner,  Feb.  16,  1905,  while  making  a  journey  up  the  Nile  river  into 
the  Sudan.  All  data  are  taken  from  Klaptocz 's  (1906)  paper  since  it  was 
impossible  to  secure  any  material  for  the  writer's  examination. 

Measurements  of  the  worm  are  as  follows :  Total  length  28.0  mm.  ; 
lateral  diameter  of  scolex  0.975  mm.,  dorso-ventral  diameter  0.83  mm. ; 
maximum  diameter  of  suckers  0.5  mm.;  diameter  of  fifth  sucker  0.085- 
0.10  mm. ;  thickness  of  proglottids  near  scolex  0.46  mm.,  breadth  0.87 
mm.,  length  0.04  mm.;  maximum  breadth  of  proglottid  near  middle 
of  the  strobila  1.19  mm.,  length  0.21  mm.;  breadth  of  sixth  proglottid 
from  end  0.59  mm.,  length  0.21  mm. ;  breadth  of  next  to  last  proglottid 
0.41  mm.,  length  0.31  mm. ;  breadth  of  last  proglottid  0.33  mm.,  length 
0.51  mm.  The  scolex  (Figs.  126,  127)  is  somewhat  oval  in  transection, 
surpassing  the  breadth  of  the  first  proglottids.  It  possesses  no  rostellum, 
no  armature  of  hooks,  but  has  an  apical  sucker  0.085-0.100  mm.  in 
diameter.  The  four  equal  sized  suckers  of  somewhat  oval  or  round  shape 
vary  in  size  in  different  individuals.     They  are  about  0.50  mm.  in 


166  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [166 

diameter  and  are  situated  on  the  four  quadrants  of  the  head.  The  four 
quadrants  are  somewhat  swollen  and  are  separated  from  each  other  by 
longitudinal  furrows.  The  apical  depression  is  a  true  sucker,  as  is  shown 
in  the  stained  specimen.  There  is  no  neck,  the  first  segments  appearing 
under  the  microscope  0.060  mm.  behind  the  margin  of  the  suckers.  The 
size  relations  of  the  proglottids  have  already  been  given.  From  the 
drawing,  reproduced  (Figs.  128,  129),  the  strobilation  is  evident,  the 
posterior  angles  of  the  proglottids  being  quite  prominent. 

The  anlagen  of  the  sexual  organs  may  be  seen  7  mm.  back  of  the 
tip  of  the  scolex  while  the  well  developed  testes  may  be  seen  3  mm.  farther 
posteriad.  These  are  elliptical  in  shape,  their  long  axis  lying  in  the 
long  axis  of  the  worm.  They  number  70-100,  and  lie  irregularly  scat- 
tered between  the  vitellaria  and  anterior  to  the  ovaries.  They  are  most 
numerous  in  the  lateral  fields  but  they  may  also  occur  in  the  median 
part  especially  in  the  anterior  region.  No  data  are  given  concerning  the 
cirrus-pouch  and  the  vas  deferens.  The  genital  pore  is  irregularly 
alternating.  It  is  situated  near  the  middle  of  the  margin  of  the  seg- 
ment, but  in  the  last  proglottids  a  trifle  anterior  thereto. 

The  vagina  opens  posterior  to  the  cirrus-pouch.  Just  before  its 
opening  it  possesses  a  considerable  broadening,  the  diameter  of  which 
is  about  equal  to  the  broadest  diameter  of  the  cirrus-pouch.  On  either 
side  of  this  broadened  part  the  vagina  is  narrow.  The  vitellaria  are 
lateral  longitudinal  glands  which  develop  late.  They  are  not  volum- 
inous. The  bi-lobed  ovary  lies  in  the  posterior  part  of  the  proglottid  as 
in  Proteocephalids.  Since  the  uterus  was  inunature  in  this  species  the 
eggs  could  not  be  observed. 

This  species  easily  separates  itself  from  the  most  of  the  forms, 
parasitic  in  fish,  by  reason  of  the  large  size  of  its  head  and  suckers. 
The  position  of  the  vagina  posterior  to  the  cirrus-pouch  is  a  further 
diagnostic  character  of  value,  while  the  lack  of  an  unsegmented  neck 
and  the  proportions  of  the  proglottids  are  characters  which  serve  for 
its  identification.  Unfortunately  Klaptocz  failed  to  describe  and  to  figure 
the  cirrus-pouch,  cirrus,  and  vas  deferens  which  are  of  great  value  in 
making  a  positive  determination.  The  uterus  being  immature  could  not 
be  described.  For  these  reasons  this  species  which  probably  is  a  species 
of  Proteocephalus  cannot  be  accurately  placed  in  its  relation  to  the 
other  members  of  the  genus. 


167]  PROTEOCEPHALIDAE—LA  RUE  167 

PROTEOCEPHALUS  FOSSATUS  (Riggenbach) 
[Figs.  133,  180] 

1896:    Ichthyotaenia  fossata  Riggenbach        1896:166-193 

1911 :    Proteocephalus  fossatus        La  Rue  1911 :475 

Specific  Diagnosis:  The  characters  of  the  genus.  Cestodes  of 
short  length,  3.5-4  cm.  long.  Scolex  large,  visible  to  nailed  eye,  maximum 
breadtli  of  same  0.714  mm.  Anterior  end  of  scolex  conical,  bearing  at 
apex  a  slight  concavity,  not  a  fifth  suclier.  Sucliers,  round,  0.34  mm.  in 
diameter,  borne  at  broadest  part  of  head.  Neck  broad,  0.85  mm.  long. 
First  proglottids  broader  than  long,  0.612  mm.  broad  by  0.135  mm.  long. 
Mature  and  ripe  proglottids,  quadrate  to  longer  than  broad,  End- 
proglottids  not  observed.  Lateral  margins  of  proglottids  quite  straight^ 
about  the  genital  sinus  a  slight  elevation  or  genital  papilla. 

Genital  aperture  marginal,  irregularly  alternating,  situated  a  little 
anterior  to  the  middle  of  the  proglottid.  Testes,  numerous,  120-150,  in 
medullary  parenchyma  between  vitellaria.  Vas  deferens,  a  loose  mass 
of  coils  between  cirrus-pouch  and  middle  of  proglottid.  Ductus  ejacu- 
latorius  with  few  coils.  Cirrus  short  and  thicl^.  Cirrus-pouch  pear- 
shaped,  0.30  mm.  long,  extending  about  y^  across  the  proglottid  breadth. 
Vagina,  opening  anterior  to  cirrus-pouch.  Vaginal  sphincter  small, 
near  opening.  Lumen  of  first  part  of  vagina  broad,  nearly  as  large  as 
cirrus-pouch.  No  coils  in  vagina  anterior  to  ovary.  Arrangement  of 
organs  in  interovarial  space  typical  of  genus.  Ovary  bilobed,  posterior. 
Vitelline  glands  follicular,  lateral,  extending  full  length  of  segment. 
Uterus  in  ripe  proglottid  with  many  lateral  outpocketings  on  either  side. 
Eggs,  round,  thin  shelled  structures,  size  not  given. 

Habitat :  Intestine  of  Pimelodus  pati  Valenc,  Rio  Paraguay,  South 
America. 

This  species  was  described  by  Riggenbach  (1896)  who  made  it  a 
member  of  the  genus  Ichthyotaenia.  La  Rue  (1911)  listed  this  species 
in  a  list  of  species  of  Proteocephalus.  The  material  was  collected  by  Dr. 
Ternetz  in  January  and  February  1894.  The  following  description  is 
based  on  Riggenbach 's  (1896)  paper. 

The  worms  as  collected  were  of  small  size,  3.5-4.0  cms.  long.  There 
was  no  undamaged  strobila.  The  neck  is  0.85  mm.  long  and  is  quite 
broad.  It  passes  imperceptibly  into  the  youngest  proglottids.  The 
scolex  (Fig.  133)  is  relatively  large,  and  is  perceptible  to  the  naked  eye. 
It  bears  four  large  suckers  at  its  broadest  zone.  Here  the  scolex  is  0.714 
mm.  broad.     From  this  point  the  scolex  becomes  smaller  toward  the 


168  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [168 

end  and  also  toward  the  neck.  Its  anterior  region  is  conical.  At  the 
apex  of  the  head  is  a  slight  depression  which  structurally  is  not  a  fifth 
sucker.  There  is  no  rostellum.  The  suckers  are  round  in  outline  and 
they  measure  0.34  mm.  in  diameter. 

The  youngest  proglottids  in  the  proliferation  zone  are  narrow  trans- 
verse bands  0.135  mm.  long  by  0.612  mm.  broad.  With  increasing  age 
the  proglottids  elongate  until  mature  and  ripe  proglottids  may  be  quad- 
rate and  the  oldest  proglottids  even  longer  than  broad.  An  end-proglot- 
tid  was  not  seen.  The  margins  of  the  segments  are  nearly  straight.  At 
the  place  of  the  opening  of  the  genital  passages  the  surface  is  slightly 
raised  to  form  a  small  genital  papilla. 

In  the  head  the  excretory  vessels  run  close  together.  Near  the 
suckers  a  circular  anastomosis  unites  the  vessels.  In  the  space  between 
the  suckers  is  a  complicated  plexus.  The  two  pairs  of  main  lateral 
excretory  vessels  run  through  the  length  of  the  worm  in  nearly  a  straight 
course.  In  the  posterior  margin  of  each  proglottid  is  a  transverse  anas- 
tomosis connecting  the  main  vessels.  In  this  region  also  and  in  the 
neck  short  ducts  arise  from  the  main  ventral  vessels  only  and  pass  to 
the  exterior,  opening  on  the  surface  near  the  posterior  angle  of  the 
proglottid.  The  lumen  of  such  ducts  becomes  smaller  near  the  periph- 
ery of  the  work.    The  passage  through  the  cuticula  is  quite  small. 

The  genital  aperture  is  marginal,  irregularly  alternating,  situated 
a  little  anterior  to  the  middle  of  the  proglottid.  A  very  small  genital 
papilla  is  present,  due  largely  to  a  thickening  of  the  margin  of  the 
sinus.  The  anlagen  of  the  genital  organs  appear  in  about  the  twentieth 
segment.  The  testes  (Fig.  180)  number  120-150  and  are  relatively 
small,  0.5  mm.  in  diameter,  round  or  polygonal  in  outline.  They  fill 
the  whole  field  between  the  two  vitelline  glands  and  the  anterior  and 
posterior  margins  of  the  proglottid.  The  vas  deferens  forms  a  loose 
mass  of  coils  between  the  cirrus-pouch  and  the  middle  of  the  proglottid. 
The  mass  lies  posteriad  to  the  cirrus-pouch  and  is  excentric  to  it,  lying 
on  the  porose  side  of  the  proglottid.  In  the  cirrus-pouch  the  ductus 
ejaculatorius  forms  a  few  coils  and  then  it  passes  over  into  the  cirrus. 
The  cirrus  is  enlarged  into  a  thick  vesicle  which  fills  up  nearly  the 
outer  half  of  the  cirrus-pouch.  The  cirrus-pouch,  which  is  about  0.30 
mm.  long,  reaches  about  J^  across  the  proglottid  breadth.  It  is  an 
elongated  oval  in  shape. 

The  vagina  which  always  opens  anterior  to  the  cirrus-pouch  has  a 
small  sphincter  near  its  opening.  Further  within,  the  lumen  of  the 
vagina  enlarges  into  an  ovoidal  vesicle  which  may  be  almost  as  volumi- 
nous and  as  long  as  the  cirrus-pouch.  The  vagina  forms  no  coils  ante- 
rior to  the  ovary.    Riggenbach  found  no  receptaculum  seminis  but  he 


169]  PROTEOCEPHALIDAE—LA  RUE  169 

found  a  widening  of  the  vagina  anterior  to  the  ovary.  The  two  dilata- 
tions of  the  vagina  take  the  place  of  the  receptaculum  seminis.  The 
arrangement  of  the  organs  in  the  interovarial  space  is  typical  of  the 
genus.  The  ovary  is  a  bilobed  structure,  situated  in  the  posterior  part 
of  the  proglottid.  The  ovary  does  not  seem  to  be  made  up  of  blind- 
pouches,  as  is  true  in  the  ovaries  of  most  of  the  Taenias.  Its  outline  is 
somewhat  irregular,  due  to  the  sack-like  processes  which  may  be  seen 
on  the  plump  mass  of  the  lobe.  These  extend  posteriad  nearly  to  the 
proglottid  margin  and  laterally  to  the  excretory  vessels.  The  vitellaria 
as  in  all  Proteocephalids  are  made  up  of  a  large  number  of  single  folli- 
cles arranged  in  long  bands  extending  the  full  length  of  each  lateral 
field  of  proglottid.  The  vitelline  ducts  arise  in  the  posterior  region  of 
the  segment  and  their  course  to  the  interovarial  space  is  like  that  of 
other  Proteocephalids.  The  shell-gland  is  poorly  developed.  The  ute- 
rus lies  in  the  middle  of  the  proglottid  and  "As  a  canal  with  numerous 
lateral  branches  it  extends  itself  through  the  whole  proglottid."  "The 
uterine  eggs  are  round  thin  shelled  structures."  Their  size  was  not 
given. 

This  species  is  readily  separated  from  a  large  number  of  species  of 
Proteocephalus  on  account  of  its  lack  of  a  fifth  sucker.  Among  those 
species  which  do  not  possess  fifth  suckers  this  species  is  most  like  P. 
sulcatus  which  likewise  occurs  in  one  of  the  Siluridae.  The  latter  spe- 
cies however  is  much  larger  as  to  head  and  as  to  observed  length.  It 
also  has  a  much  larger  number  of  testes.  The  form  of  the  head  is  also 
different. 


PROTEOCEPHALUS  SKORIKOWI  (Von  Linstow) 
[Figs.  152,  177] 

1904:    Ichthyotaenia  skorikowi  von  Linstow      1904:18-19 

1911:     Proteocephalus  skorikowi         La  Rue  1911:475 

This  species  was  described  by  von  Linstow  (1904).  La  Rue  (1911) 
listed  it  among  other  species  of  Proteocephalus.  The  following  descrip- 
tion is  an  abstract  from  von  Linstow 's  account. 

The  length  measures  as  much  as  200  mm.,  the  breadth  anteriorly 
1.14  mm.  First  proglottids  beginning  directly  behind  the  scolex  are 
very  short.  Breadth  in  middle  is  3.75  mm;  the  length  of  proglottids 
here  0.95  mm.  Breadth  at  posterior  end  equals  3.16  ram;  length  of 
proglottids  here  1.97  mm.  The  last  proglottid  is  rounded  posteriorly. 
The  scolex  is  short,  0.67  mm.  broad.  Von  Linstow 's  drawing  of  the  head 
is  reproduced    (Fig.   152).     Suckers  measure  0.250  mm.  in  diameter 


170  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [170 

while  a  fifth  apical  sucker  is  0,130  mm.  in  diameter.  Muscles  of  sub- 
cuticular layer  are  as  usual.  The  heavy  layer  of  longitudinal  muscles 
below  the  subcuticula  in  the  anterior  part  of  the  worm  is  very  strongly 
developed,  filling  up  almost  the  entire  space  in  unripe  proglottids. 
Dorsoventral  muscles  are  very  heavily  developed.  In  anterior  segments 
the  main  longitudinal  nerves  lie  about  ^Vioo>  and  in  ripe  segments  about 
'/ioo>  of  the  segment  width  from  the  lateral  margins.  Mesad  of  the 
nerve  trunks  lie  the  excretory  ducts  two  on  either  side.  Chalkbodies 
are  lacking  entirely. 

Sexual  openings  are  marginal,  irregularly  alternating,  posterior  to 
the  middle  in  each  proglottid.  The  vagina  lies  anterior  to  the  cirrus. 
The  cirrus  is  cone-shaped,  blunt  at  the  tip,  0.141  mm.  long.  The  point 
is  0.042  mm.  broad.  The  cirrus-sheath  extends  Vq-Yq  across  the  breadth 
of  the  proglottid.  The  vas  deferens  makes  many  coils  before  entering 
the  cirrus-sheath.  Testes  are  very  numerous,  0.053-0.083  mm.  in  diam- 
eter. From  von  Linstow's  figure  one  may  conclude  that  there  are  about 
70-100  testes.  The  vagina  runs  in  a  curve  to  the  middle  of  the  posterior 
region  of  the  segment  where  it  forms  many  coils  anterior  to  the  ovaries. 
These  coils  function  as  a  receptaculum  seminis.  The  ovary  is  made  up 
of  two  groups  of  club-shaped  bags  which  run  parallel  to  the  posterior 
margin  of  the  proglottid.  Anterior  to  its  middle  lies  an  oocapt.  Vitel- 
laria  are  follicular,  reaching  on  left  and  right  the  full  length  of  the 
proglottid  inside  the  longitudinal  muscles.  The  uterus  in  the  posterior 
part  fills  the  whole  space  within  the  longitudinal  muscle  fibers  and 
extends  anteriad  in  round  diverticula.  In  von  Linstow's  drawing  which 
is  reproduced  (Fig.  177)  there  are  6-8  diverticula  on  either  side.  Eggs 
are  globular,  0.027  mm.  They  have  two  membranes,  the  outer  one  very 
delicate  and  hyaline.  The  six-hooked  embryo  measures  0.021-0.023  mm. 
in  diameter. 

Habitat:  Intestine  of  Acipenser  steUatus  (t3T)e  host);  River 
Giirgen,  Caspian  Sea  (type  locality).  Von  Linstow  failed  to  state  who 
collected  the  specimens  of  this  species  and  the  abundance  of  their 
occurrence. 

This  species  is  one  of  the  more  robust  Proteocephalids  and  is  thus 
distinguished  from  a  large  number  of  species  of  the  genus.  The  fact 
that  the  vagina  forms  many  coils  anterior  to  the  ovary  and  that  the 
ovary  consists  of  slender  club-shaped  bags  separates  it  effectively  from 
all  other  known  species.  The  position  of  the  genital  pore  posterior  to 
the  middle  of  the  proglottid  is  a  further  distinguishing  feature.  In 
many  respects  this  species  most  closely  resembles  P.  fossatus  (Riggen- 
bach). 


171]  PROTEOCEPHALIDAE—LA  RUE  171 


PROTEOCEPHALUS  SAGITTUS  (Grimm) 


(?)1819: 

Bothriocephalus  barbatulae 

Rudolphi 

1819 :144 

(?)1850: 

Bothriocephalus  harhatulae 

Diesing 

1850 :608 

1872: 

Taenia  sagitta 

Grimm 

1872 :240-243 

1878: 

Taenia  sagittata 

von  Linstow 

1878 :260 

1891: 

Taenia  sagittata 

Monticelli 

1891 :169 

1896: 

Ichthyotaenia  sagittata 

Riggenbach 

1896 :267 

1909: 

Ichthyotaenia  sagitta 

Liihe 

1909 :33 

1911: 

Proteocephalus  sagittus 

La  Rue 

1911 :475 

Specific  Diagnosis:  Strobila  small,  as  much  as  45  mm.  long  by  1 
mm.  broad.  Head  thick,  ending  in  an  unarmed  point.  Suckers  four, 
in  pairs  on  the  basal  part  of  the  head.  Suckers  transversely  elongated 
and  heavily  muscled.  Neck  as  much  as  12  mm.  long  by  0.5  mm.  broad. 
(Probably  many  young  proglottids  were  overlooked  in  this  exception- 
ally long  neck.)  Proglottids  few.  Youngest  proglottids  about  quad- 
rate, older  ones  longer.  Last  eight  proglottids  1.4-1.5  mm.  long  by  1 
mm.  broad  by  0.3  mm.  thick.  Last  proglottid  always  rounded  poste- 
riorly. Cirrus-pouch  large  and  clubshaped.  Cirrus  quite  thick.  Uterus 
with  uterine  pouches,  number  unknown.  Genital  pore  situated  near 
middle  of  lateral  margin  of  proglottid. 

Habitat:  In  intestine  of  Cdbitis  harbatula  (type  host),  Peters- 
burg (type  locality)  and  Province  of  Novgorod. 

Grimm  (1872:240-243)  described  some  cestodes  which  he  had  col- 
lected from  the  intestines  of  Cohitis  harhatula  at  Petersburg  and  in  the 
province  of  Novgorod.  These  he  named  Taenia  sagitta.  He  called  at- 
tention to  the  fact  that  Rudolphi  (1819:144)  had  proposed  the  name 
Bothriocephalus  harhatulae  for  some  cestodes  from  Cohitis  harhatula. 
Rudolphi  gave  no  diagnosis  nor  description  hence  it  is  not  possible  to 
determine  the  genus  to  which  his  specimens  belonged.  Grimm  also 
called  attention  to  Diesing's  (1850:608)  reference  to  Bothriocephalus 
harhatulae  and  that  Diesing  gave  no  description  of  the  form.  Von 
Linstow  (1878:260)  listed  Taenia  sagittata  Grimm  (for  sagitta)  as  a 
parasite  of  Cohitis  harhatula.  Monticelli  (1891:169)  included  Taenia 
sagittata  (for  sagitta)  in  a  list  of  cestode  species  parasitic  in  fish.  Rig- 
genbach (1896:267)  included  Ichthyotaenia  sagittata  (Grimm)  in  a  list 
of  Ichthyotaenia.  Liihe  (1909:33)  gave  a  very  short  diagnosis  of  Ich- 
thyotaenia sagitta  (Grimm)  which  he  apparently  derived  from  Grimm's 
(1872)  paper.  La  Rue  (1911:475)  included  this  species  in  a  list  of 
Proteocephalus  species. 


172  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [172 

Because  of  its  inaccessibility  to  many  scientific  workers  Grimm's 
entire  description  of  this  species  is  here  quoted,  and  since  the  tables  at 
the  end  of  this  section  give  the  essential  data  no  digest  of  his  paper  is 
given  at  the  end  of  his  description : 

\J  "TAENIA  SAGITTA. — Der   beriihmte    Helmintholog    Rudolphi    erwahnt    in 

seinem  Werk  Entozoorum  Synopsis  S.  144  eines  Bandwurmes,  den  er  im  Darme 
von  Cobitis  barbatula  aufgefunden  hat.  Da  er  aber  nur  ein  Fragment,  nicht  aber 
ein  voiles  Exemplar  des  Wurmes  besass,  so  gab  er  auch  keine  eingehendere  Be- 
schreibung  dieser  als  neu  von  ihm  erkannten  Art,  sondern  stellte  sie  grade  zu  den 
Grubenkopfen  und  belegte  sie  mit  dem  Namen  Bothriocephalus  barbatulae.  Dies- 
ing  stellte  diese  Art  in  seinem  Systema  helminthum  zu  den  Species  dubiae  und 
setzte  zu  den  von  Rudolphi  gesagten  nur  noch  hinzu,  dass  "Fragmenta  acephala 
servantur  im  M.  C.  V."  (Syst.  helm.  I.  p.  608).  Etwas  umstandlicher  spricht  sich 
S.  Leuckart  aus  iiber  das  Aussehen  dieses  Helminthen,  indem  er  sagt,  "an  dem  sehr 
kleinen  kopflosen  Stiicke,  das  ich  sah,  war  nichts  Ausgezeichnetes.  Die  Glieder 
langer  als  breit,  oder  vollkommen  quadratisch.  Keine  deutlichen  Ovarien"  (Sig. 
Leuckart,  Zoologische  Bruchstiicke.  I.  p.  57).  Das  ist  Alles,  was  wir  von  diesem 
Bandwurme  kennen  und  deshalb  wird  wohl  nicht  uberflussig  sein,  wenn  ich  im  fol- 
genden  die  Resultate  meiner  anatomischen  Untersuchung  kurz  zusammen  fassen 
werde,  die  ich  an  diesem  Wurm  unlangst  angestellt  habe. 

"Ich  fand  namlich  hier,  in  Petersburg,  und  im  Nowgorod'schen  Gouverne- 
ment  3  erwachsene  Exemplare  des  Wurms  in  Cobitis  barbatula,  nachdem  ich  einige 
Hunderte  dieses  so  haufig  in  unseren  kleineren  Flussen  vorkommenden  Fischchens 
durchmustert  hatte.  Aus  dem  Gesagten  geht  schon  hervor,  dass  dies  ein  sehr 
seltner  Wurm  sein  muss.  Hier  muss  ich  noch  bemerken,  dass,  dem  Kopfe  nach, 
dieser  Wurm  der  Taenia  filicollis  Rud.  nicht  unahnlich  ist,  sich  aber  von  ihr  durch 
die  langeren  Glieder,  wie  wir  es  unten  sehen  werden,  unterscheidet. 

"Die  Lange  des  grossten  Exemplares  betrug  45  Mm.  bei  einer  Breite  von  i 
Mm.  Der  Kopf  ist  verdickt  und  endigt  mit  einer  Spitze  die  keine  Hacken  tragt; 
die  4  Saugnapfe  liegen  paarweise  auf  der  Grundhalfte  des  Kopfes;  sie  sind  etwas 
in  die  Quere  verlangert  und  stark  muskulos,  weshalb  der  Wurm  auch  so  fest  an 
die  Wande  des  Darms  sich  ansaugt,  dass  man  beim  Einsammeln  sehr  vorsichtig 
sein  muss,  wenn  man  nicht  "Fragmenta  acephala"  erhalten  will.  Das  zu  seiner 
Basis  verengte  Kopfchen  geht  in  den  ziemlich  diinnen  Hals  tiber,  welcher  ungefahr 
12  Mm.  lang  und  0,5  Mm.  breit  ist.  Hinter  dem  Halse  fangt  die  Kette  an,  in  der 
ich  23  einzelne  Glieder  gezahlt  habe.  Die  ersten  resp.  jungsten  Glieder  sind  fast 
vollkommen  quadratisch ;  weiterhin  werden  sie  etwas  langer,  so  dass  die  letzten 
acht  Glieder  14  bis  1,5  Mm.  lang  sind;  dabei  betragt  ihre  Breite  i  Mm.,  bei  einer 
Dicke  von  nur  1,3  Mm.,  so  dass  der  Wurm  vollkommen  plattgedriickt  erscheint. 
Das  letzte  Glied  ist  immer  von  hinten  abgerundet  Dies  ist  alles  was  wir  an 
einem  unladirten  Exemplare  zu  sehen  bekamen. 

"Wenn  wir  nun  aber  einzelne  zuvor  mit  Carmin  und  Glycerin  bearbeitete 
Glieder  und  diinne  Querschnitte  unter  dem  Mikroskop  untersuchen,  so  iiberzeugen 
wir  uns,  dass  der  Bau  der  Korpermasse  dieses  Bandwurmes  sich  iiberhaupt  durch 


173]  PROTEOCEPHALIDAE—LA  RUE  173 

nichts  von  dem  Bau  anderer  Bandwiirmer  unterscheidet.  Wir  finden  hier  wie 
auch  bei  Cyathocephalus  truncatus,  dass  der  Korper  aus  folgenden  Elementen 
besteht : — aus  der  kornchenreichen  Grundsubstanz,  die  von  sich  schlangelnden 
dorso-ventralen  Muskelbundeln  durchsetzt  wird;  einer  Schicht  Langen-muskeln 
und  einer  Schicht  Quer — oder  Ringmuskeln,  einer  Schicht  feinkorniger  Rinden- 
substanz,  in  der  die  Enden  der  dorso-ventralen  Muskeln  liegen,  und  endlich  der 
sehr  feiner  ausseren  Haut,  die  gar  keine  Muskeln  besitzt.  In  der  Langsaxe  eines 
jeden  reifen  Gliedes  liegt  das  centrale  Rohr  des  Eierstocks,  welcher  aber  manch- 
mal  naher  zur  einen  oder  andern  flachen  Seite  zu  liegen  kommt,  wie  man  es  auf 
den  Querschnitten  bemerkt.  Von  diesem  Centralrohr  verlaufen  nach  beiden  Seiten 
zu  unregelmassig  gebogene  und  theils  sich  auch  verzweigende  Auslaufer,  die  in 
ihrer  Lange  mittelst  der  dorso-ventralen  Muskeln  erhalten  werden.  In  der  Mitte 
des  Hauptrohres  geht  von  ihm  die  Scheide  ab,  die,  nachdem  sie  eine  schwache 
Biegung  gemacht  hat,  sich  in  der  Mitte  der  seitlichen  Oberfliiche  des  Gliedes  nach 
aussen  offnet;  die  Scheide  ist  ziemlich  breit,  so  dass  die  Eier  sie  leicht  passiren 
konnen.  In  dem  von  mir  untersuchten  Exemplare  waren  die  Eier  in  der  Mehrzahl 
der  Glieder  vollkommen  reif ; — sie  enthielten  eine  grob  und — feinkornige  Masse 
und  waren  mit  einer  ziemlich  dicken  Haut  umgeben.  Aber  wie  in  diesen  reifen 
Gliedern,  so  auch  in  den  jiingern  bin  ich  nicht  im  Stande  gewesen  die  sog. 
Dotterstocke  auf zufinden ;  ob  sie  hier  so  f riih  verschwinden,  oder  zu  fein  sind, 
weiss  ich  nicht;  auf  den  Querschnitten  sind  sie  aber  nicht  aufzufinden. 

"Gleich  iiber  der  Scheide  liegt  und  offnet  sich  nach  aussen  das  mannliche 
Zeugungsorgan ;  dasselbe  besteht  aus  einem  ziemlich  grossen  und  kolbenf ormigen 
Cirrusbeutel,  in  dem  der  eingezogene  Cirrus  liegt;  der  Cirrus  ist  ziemlich  dick 
und  am  Grunde  dunner  als  am  Ende;  dessen  Kanal  ist  durch  den  Cirrusbeutel  zu 
sehen.  In  den  Wanden  des  Cirrusbeutels  trafen  wir,  wie  immer,  Langs — und 
Quermuskeln.     Die  Samendriisen  waren  nicht  zu  sehen. 

"Dies  ist  alles  was  ich  von  dem  Bau  des  Bandwurmes  erfahren  konnte.  Die 
Unvollkommenheit  meiner  Untersuchung  wurde  aber  durch  den  Mangel  an  Mate- 
rial bedingt.  Dennoch  aber  fiihle  ich  mich  berechtigt  zu  sagen,  dass  auch  dies 
Wenige  geniigt,  um  dem  beschriebenen  Bandwurm  seine  natiirliche  Stellung  in  der 
Reihe  seines  Gleichen  anzuweisen  und  somit  die  Zahl  der  zweifelhaften  Arten  zu 
beschranken. 

"Noch  muss  ich  hinzusetzen,  dass  aus  dem  Gesagten  doch  klar  ist,  dass  ich 
wohl  mich  nicht  irre,  indem  ich  den  von  mir  augefundenen  Wurm  als  dem  Both- 
riocephalus  barbatulae  Rud.  identisch  halte." 

While  the  foregoing  description  is  sufficient  to  warrant  the  inclu- 
sion of  this  species  in  the  genus  Proteocephalus  it  is  not  sufficient  to 
enab.e  one  to  determine  the  relations  of  the  species  to  the  other  mem- 
bers of  the  genus.  In  size  the  form  resembles  P.  filicollis  and  P.  agonis. 
For  the  present  the  species  must  be  put  in  the  list  of  incompletely 
described  species. 


174  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [174 

PROTEOCEPHALUS  SALVELINI  (Linton) 
[Figs.  143,  144] 
1897:     Taenia  salvelini  Linton  1897:424 

1911:    Proteocephalus  salvelini       La  Rue  1911:475 

Linton  (1897:424)  described  this  species.  La  Rue  (1911:475)  in- 
cluded this  form  in  a  list  of  species  of  Proteocephalus. 

Linton's  description  reads: 

"TAENIA  SALVELINI.  Type.—l<io.  481 1,  U.  S.  N.  M.  From  intestine  of 
Great  Lake  trout  (Cristivoiiier  namaycush)  ;  Outer  Island,  Lake  Superior;  J.  W. 
Milner.  In  this  lot  are  several  small  Taeniae;  scolices  with  few  roundish  or 
oblong  segments,  in  some  of  which  the  male  genitalia  were  developed,  otherwise 
immature;  length  from  3  to  5  mm. 

"These  specimens  bear  some  resemblance  to  the  young  of  T.  torulosa,  Batsch, 
as  described  by  Zschokke   (1884). 

"Head  usually  rounded  in  front,  with  suckers  directed  anteriorly.  The  diam- 
eter of  the  head  varies  according  to  state  of  contraction,  usually  0.3  to  0.4  mm.; 
suckers  relatively  large,  usually  directed  forward,  prominent,  rather  deep.  0.15  to 
0.18  mm.  in  diameter,  aperture  o.i  mm.  in  diameter.  Neck  assuming  various  con- 
traction shapes ;  in  some  continuous  with  head  and  merging  imperceptibly  in  body ; 
in  others  separated  by  a  constriction  behind  the  head;  in  others  not  separated 
from  head  by  constriction  but  with  segments  beginning  abruptly;  numerous  cal- 
careous bodies  in  neck  behind  head.  Usually  first  segments  indistinct,  broader 
than  long.  Oldest  segments  longer  than  broad;  genital  aperture  near  middle  of 
lateral  margin  a  little  nearer  anterior  margin.  Vitellaria  making  a  narrow  border 
along  lateral  margins ;  beginning  of  germ  gland  at  posterior  end ;  central  region 
in  front  of  rudimentary  germ  gland  occupied  by  prominent  testicular  lobes ;  cirrus- 
bulb  oblong,  elliptical,  convex  on  posterior,  straight  or  slightly  concave  on  anterior 
surface;  cirrus  enters  from  inner  anterior  edge  of  bulb,  then  traverses  middle  of 
bulb  to  orifice ;  length  of  bulb,  0.26  mm. ;  diameter,  0.14  mm. ;  largest  segments, 
1.5  mm.  in  length  and  0.75  mm.  in  breadth." 

Linton's  figures  of  the  head  are  reproduced  (Figs.  143,  144). 

Since  this  description  is  insufficient  for  determination  in  compari- 
son with  the  forms  described  here  Dr.  H.  B.  Ward  very  kindly  secured 
Linton's  specimens  from  the  National  Museum  at  Washington.  Unfor- 
tunately but  fragments  remained.  These  fragments  were  cleared  and 
examined  in  glycerine.  The  writer's  observations  on  the  heads  agree 
very  well  with  Linton's  records.  A  fifth  sucker  or  a  trace  of  one  could 
not  be  found.  The  suckers  in  their  maximum  diameter  measure  0.17O 
mm.  the  sucker  opening  about  0.100  mm.  The  head  is  flattened  dorso- 
ventrally.  In  general  shape  it  resembles  some  dilated  heads  of  P.  pin- 
guis  but  is  larger.  Linton  says  that  it  resembles  young  specimens  of 
P.  tornlosus,  however  its  measurements  are  quite  different.  It  must  be 
left  for  the  present  in  the  list  of  inadequately  described  species. 


175]  PROTEOCEPHALIDAE—LA  RUE  175 

PROTEOCEPHALUS  CYCLOPS  (Von  Linstow)  sp.  inq. 

[Fig.  159] 
1877 :     Taenia  cyclops  von  Linstow,  1877 :15-16. 

Von  Linstow 's  description  of  this  immature  form  is  here  given: 
"Taenia  cyclops  n.  sp. — In  Coregonus  maraena  aus  dem  Schallsee.  Die  Tanie 
ist  25  Mm.  lang  und  0,27  Mm.  breit,  iiberall  fast  gleich  breit,  die  letzte  Proglottide 
hinten  zugespitzt.  Die  Kalkkorperchen  sind  klein,  ohne  concentrische  Schichtung; 
das  Kopfende  ist  abgerundet,  die  Saugnapfe  sind  langsoval,  0,15  Mm.  lang  und 
nach  hinten  zugespitzt,  0,1  Mm.  breit;  ein  funfter  scheitelstandiger  Saugnapf  hat 
0,069  Mm.  im  Durchmesser;  Geschlechtsorgane  waren  noch  nicht  vorhanden. 
Taenia  longicollis  aus  verschiedenen  zur  Familie  der  Lachse  gehorenden  Fischen 
hat  zum  Unterschied  von  dieser  Art  langlichrunde  Saugnapfe,  deren  langerer 
Durchmesser  rechtwinklig  zur  Langsaxe  des  Thieres  steht." 

Nufer  (1905:152)  thought  that  this  form  was  a  lariFat -state  of  P. 
longicollis  (Rud.).  However,  the  character  of  the  sucJiers  does  not  per- 
mit such  a  determination.  P.  cyclops  is  remarkable  for  the  notch  in 
the  inferior  margin  of  the  sucker  and  in  this  respect  differs  from  all 
the  other  species  of  Proteocephalus.  This  notch  is  well  shown  in  von 
Linstow 's  drawing  which  is  reproduced  (Fig.  159).  Until  adults  can 
be  secured  for  determination  this  form  must  be  considered  to  be  a  spe- 
cies inquirenda. 


PROTEOCEPHALUS    HEMISPHERICUS   (MoUn)   sp.  inq. 


1859 
1896 
1911 


Taenia  hemispherica  Molin  1859:14 

Ichthyotaenia  hemisphaerica    Riggenbach    1896:267,  268 
Proteocephalus  hemisphericus  La  Rue  1911 :475 


Molin 's  original  description  and  observations  are  as  follows: 

"Taenia  hemispherica  Molin. — Caput  hemisphericum,  acetabulis  anticis;  ore 
inermi;  collum  longissimum,  antice  dilatatum;  articuli  supremi  brevissimi,  subse- 
quentes  transverse  parallelogrammici,  postremi  subquadrati ;  aperturae  genitales 
marginales,  vage  alternae.    Longit.  0.07;  lat.  0.003. 

"HABITACULUM.  Anguilla  vulgaris:  in  intestino  tenui,  Novembri,  Patavii 
(Molin). 

"OSSERVAZIONE  i.  Ai  5  di  Novembre  del  1858  rim-enni  nell'  intestino  di 
un'  anguilla,  nella  quale  avevo  trovato  5  Dibothrium  claviceps,  un  esemplare  di 
quel  verme ;  ed  in  un'  altra  anguilla  un  esemplare  intero  e  varii  f rammenti. 

"OSSERVAZIONE  2.  Negli  articoli  bene  sviluppati  potei  distinguere  ai 
margini  laterali  le  ovaja,  e  tutto  il  resto  dell'  articolo  riempito  dall'  amplissimo 


176  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [176 

ovidotto  rav\-X)lto  in  ambagi  e  ripieno  di  uova  sferiche.  Nel  mezzo  vidi  collocato 
orizontalmente  il  membro  virile  attortigliato  che  si  estendeva  fino  al  centre  dell' 
articolo. 

"OSSERVAZIONE  3.  Fino  ad  ora  non  si  conosceva  dell'  anguilla  altro  che 
la  Taenia  macrocephala,  dalla  quale  pero  la  hemisphaerica  si  distingue  per  la  forma 
della  testa  non  che  per  la  lunghezza  del  collo.  Questa  e  affine  alia  filicollis,  dalla 
quale  pero  si  distingue  per  la  forma  della  testa  e  del  collo,  e  deve  percio  venir 
registrata  dopo  di  questa  nel  sistema." 

Since  the  appearance  of  Molin's  paper  no  further  attempts  have 
been  made  to  describe  this  species.  There  is  in  fact  no  further  notice 
of  the  finding  of  this  species.  Linton  (1886)  thought  that  his  Taenia 
dilatata  might  be  the  same  as  Taenia  hemispherica.  Riggenbach  (1896) 
said  of  it,  "Ein  genauer  Vergleich  der  Diagnosen,  welche  die  genannten 
Autoren  den  fraglichen  Species  gaben,  zwingt  mich  mit  Bestimmtheit 
eine  Identitat  der  I.  (Ichthyotaenia)  dilatata  Linton  mit  I.  hemispher- 
ica Molin  anzunehmen."  Linton's  species  has  been  determined  by  the 
writer  to  be  identical  with  Proteocephalus  macrocephalus  (Creplin). 
Schneider  (1903)  identified  a  cestode  taken  from  Anguilla  vulgaris  as 
Ichthyotaenia  hemispherica.  Later  (1905)  he  decided  that  this  form 
was  not  7.  hemispherica  but  7.  macrocephala  (Creplin).  His  statement 
of  this  is  discussed  in  the  historical  summary  of  Proteocephalus  macro- 
cephalus. Nufer  (1905)  attempted  to  show  the  identity  of  P.  dilatatus 
(Linton)  and  P.  hemisphaericus  (Molin)  and  further  that  these  species 
were  identical  with  P.  macrocephalus  (Creplin).  This  contention  is 
true  in  part  at  least.  In  the  discussion  of  P.  macrocephalus  in  another 
part  of  this  monograph  identity  of  P.  dilatatus  and  P.  macrocephalus 
has  been  shown.  The  present  form,  however,  is  so  little  known  and  so 
poorly  described  that  it  seems  advisable  for  the  present  to  consider  that 
it  is  not  identical  with  the  well  known  species  of  Proteocephalus  found 
in  Anguilla  but  that  it  is  a  species  inquirenda.  La  Rue  (1911:475) 
listed  this  form  among  other  species  of  Proteocephalus. 


177]  PROTEOCEPHALIDAE—LA  RUE  177 

PEOTEOCEPHALUS   MACROPHALLUS  (Diesing)  sp.  inq. 


1850: 

Taenia  macrophalla 

Diesing 

1850:514 

1856 

Taenia  macrophalla 

Diesing 

1856:35 

1864 

Taenia  macrophalla 

Diesing 

1864 :377 

1891 

Taenia  macrophalla 

Montieelli 

1891 

1896 

Ichthyotaenia  macrophalla 

Riggenbach 

1896 :267 

This  form  is  known  only  from  the  writings  of  Diesing  listed  above. 
His  statements  are  wholly  inadequate  for  purposes  of  comparison  and 
his  drawings  (Diesing  1856,  PI.  VI,  figs.  15-20)  show  but  little  more 
than  external  features  and  the  cirrus  pouch.  It  seems  to  the  writer 
however  that  this  species  should  be  considered  as  belonging  to  the  genus 
Proteocephalus  and  that  it  is  a  species  inquirenda.  Montieelli  (1891) 
and  Riggenbach  (1896)  included  this  species  in  their  groups  of  fish 
cestodes  the  most  of  which  are  now  included  in  the  genus  Proteocepha- 
lus. 

Diesing's  original  species  description  (Diesing  1850:514)  is  here 
quoted :  ' '  Taenia  macrophalla  Diesing.  Caput  rotundatum  depressum, 
acetabulis  anticis  maximis.  Collum  nullum.  Articuli  supremi,  sub- 
quadrati,  subsequi  oblonge-quadrati,  ultimo  longissimo  rotundato.  Aper- 
turae  genitalium  marginales.  Penes  alterni  longissimi  filiformes.  Long. 
1-3'  " ;  latit.yg'  ". 

"  Habitaculum.  Cichla  monoculus,  Junio,  Julio  et  Octobri  in  Bra- 
silia (Natterer:  in  intestinis.  M.  C.  V.) 

"b.     Os  limbo  elevato,  uncinulorum    corona    interdum    decidua 
armatum. ' ' 


PROTEOCEPHALUS    NEMATOSOMA    (Leidy)   sp.  inq. 

1888:  Taenia  leptosoma  Leidy  1888:169,  (nee  Diesing) 

1891:  Taenia  nematosoma  Leidy  1891:410-418 

1891 :  Taenia  somatolepta  Montieelli  1891 

1911:  Proteocephalus  nematosoma  ha,  Une       1911:475 

In  May  1888  Leidy  reported  thus  on  the  "Parasites  of  the  Pick- 
erel". 

"Dr.  Leidy  remarked  that  among  the  numerous  parasites  which  are  men- 
tioned as  infesting  the  pike,  Esox  lucius,  of  Europe,  no  Taenia  is  indicated.  In 
the  Pickerel,  Esox  reticulatus,  brought  to  our  market,  a  species  of  the  latter  ap- 
pears to  be  common.  In  two  fishes  he  found  half  a  dozen  in  the  intestine  and 
stomach;   and   in   another  a   single  individual  two   feet  in  length.     It  resembles 


178  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [178 

closely  the  Taenia  ambloplitis,  noticed  in  the  Rock  Bass,  Amhloplites  rupestris 
(Proc.,  1887,  23),  and  may  be  the  same.  Distinguishing  it  with  the  name  of 
Taenia  leptosoma,  its  characters  are  as  follows :  Body  long  and  thin,  and  at  the 
forepart  thread-like.  Head  unarmed,  without  rostellum,  with  four  equidistant 
hemispherical  bothria;  neck  very  short  or  none;  anterior  segments  transversely 
linear,  many  times  wider  than  long;  posterior  segments  gradually  becoming  pro- 
portionately longer  and  quadrate  and  barrel  shaped;  genital  apertures  marginal, 
alternating  irregularly.    Ova  spherical. 

"Length  from  6  to  9  and  26  inches,  shortening  to  one  half  or  less ;  breadth  to 
2  and  2.5  mm.  Head  0.25  to  0.5  mm.  broad;  bothria  0.125  to  0.175  vnm.  Anterior 
segments  an  inch  from  the  head  0.175  rnrn-  long  by  I  mm.  broad;  posterior  seg- 
ments 0.5  to  0.75  mm.  long  by  2  to  2.5  mm.  broad.  Ova  0.028  to  0.032  mm.  in 
diameter. 

"A  single  slender  Scolex  associated  with  the  longest  Taenia  was  4  mm.  long 
by  0.25  wide,  but  elongated  to  8  mm.  by  o.i  wide.  The  head  was  of  the  same  form 
as  that  of  the  Taenia.  After  being  in  alcohol,  the  head  of  the  Scolex  was  0.225 
mm.  wide,  with  the  bothria  o.i  in  diameter.  The  posterior  part  of  the  body  ex- 
hibited traces  of  segmentation,  with  the  segments  0.075  rn*"-  long  by  0.25  wide." 

In  November  1890  from  the  same  host  species  Leidy  reported  Tae- 
nia nematosoma.    His  description  is  quoted: 

"Taenia  nematosoma,  n.  s. — Head  rounded  quadrate,  unarmed,  with  equi- 
distant hemispherical  bothria  and  a  small  central  papilla;  neck  short  or  none; 
fore  part  of  body  linear;  anterior  segments  much  wider  than  long;  posterior  seg- 
ments gradually  becoming  proportionately  longer,  quadrate  or  barrel-shaped;  geni- 
tal apertures  marginal  and  alternating  irregularly.  Length  to  9  inches,  contracting 
to  about  one-half.  Breadth  of  head  0.375  to  0.5  mm.;  bothria  0.175  wide;  neck 
0.25  wide;  anterior  segment  an  inch  from  the  head  0.175  mm.  long  by  2  mm, 
broad ;  posterior  segments  0.7^"  mm.  long  by  2  mm.  broad,  and  when  contracted 
widening  to  2.5  mm.     Ova  spherical,  0.028  to  0.032  mm. 

"A  half  dozen  specimens  in  the  stomach  of  two  Pickerel,  Esox  reticulatus." 

Neither  report  was  accompanied  by  drawings. 

Leidy 's  two  forms  agree  in  practically  every  particular  as  to 
measurements  and  proportions  and  host  species.  The  writer  therefore 
concludes  that  they  are  identical  and  includes  them  both  under  the 
name  P.  nematosoma  (Leidy).  This  species  differs  from  P.  ambloplitis 
chiefly  in  the  measurement  of  the  suckers.  Its  measurements  vary 
greatly  from  those  of  P.  pinguis  3nd  from  P.  esocis  Schneider.  None 
of  Leidy 's  specimens  remain,  so  the  species  must  continue  a  species 
inquirenda.  Monticelli  (1891)  on  account  of  the  confusion  of  this 
name  with  Taenia  leptosoma  Diesing  suggested  the  name  Taenia  somato- 
lepta.  However  the  name  nematosoma  should  be  preferred  because  it 
is  an  earlier  available  name  than  somatolepta.  La  Rue  (1911:475) 
made  this  one  of  his  list  of  Proteocephalus  species. 


179]  PROTEOCEPHALIDAE—LA  RUE  179 

PEOTEOCEPHALUS  SALMONIS-UMBLAE  (Monticelli)  sp.  inq. 

(?)  1811-12:    Taenia  salmonis  omul  Pallas  1811-12:409 

1884:  Taenia  salmonis  umhlae  Zschokke  1884:18-19 

1891 :  Tetracotylus  salmonis-umblae  Monticelli  1891 : 

(?)1891:  Tetracotylus  salmonis-omul  Monticelli  1891: 

1896:  Ichthyotaenia  salmonis-umhlae  Riggenbach  1896:267 

1896 :  Ichthyotaenia  salmonis  umMae  Zschokke  1896 :783 

1909 :  Ichthyotaenia  salmonis  umblae  Liihe  1909 :33 

1911 :  Proteocephalus  salmonis-umhlae  La  Rue  1911 :475 

This  species  was  first  described  and  delineated  by  Zschokke  (1884) 
who  found  it  in  Salmo  umhla,  Lake  Geneva.  His  description  and  draw- 
ings are  inadequate  for  a  determination  of  its  position.  Monticelli 
(1891)  considered  this  to  be  a  species  of  Tetracotylus.  He  also  stated 
that  he  considered  Taenia  salmonis-omul  Pallas  a  species  dubia  and 
probably  a  synonym  of  T.  salmonis-umhlae.  Monticelli  wrote  both  of 
these  names  with  a  hyphen  and  hence  was  the  first  to  give  these  species 
binary  names.  Kramer  (1892)  does  not  discuss  the  species.  Riggen- 
bach (1896)  considered  it  to  be  a  species  of  Ichthyotaenia.  He  did  not 
add  anything  to  our  knowledge  of  its  structure. 

Zschokke  (1896:783)  listed  this  species  in  his  summary  of  parasites 
found  in  the  fishes  of  Lake  Geneva.  He  did  not  find  it  in  any  of  the 
fish  from  the  Rhine  altho  he  examined  nearly  1200  fish.  Nufer  (1905) 
who  investigated  the  fish  of  Lake  Lucerne  made  no  report  on  this  spe- 
cies. Liihe  (1909)  placed  this  species  in  the  genus  Ichthyotaenia.  Since 
his  diagnosis  is  short  and  concise  it  is  here  quoted  in  lieu  of  Zschokke 's 
description  (1884:18-19): 

^^Ichthyotaenia  salmonis  umhlae  Zschokke.  30-50  cm.  lang,  1-2  mm. 
breit.  Am  Scheitel  eine  leichte  saugnapfahnliche  Vertiefung.  Proglot- 
tiden  langer  als  breit,  ca.  100  bis  150  an  Zahl,  Hodenblaschen  zahlreich ; 
ein  hodenfreies  Mittelfeld  scheint  zu  fehlen.  Cirrus-beutel  bis  fast  zur 
Mittellinie  reichend ;  Genitaloffnung  ungefahr  an  der  Grenze  von  2.  und 
3.  Fiinftel  der  Proglottidenlangs.  Uterus  mit  " zahlreichen "  (ca.  6  ?) 
Blindsacken  jederseits.  Kalkkorperchen  sehr  zahlreich,  besonders  in 
Scolex  und  Hals. 

' '  Im  Darm  von  Salmo  salvelinus  L. ;  bisher  nur  aus  dem  Genfer 
See  bekannt." 

This  species  is  too  little  known  to  permit  a  determination  of  its 
position  in  the  genus.  On  account  of  its  larger  size  and  more  numerous 
testes  it  seems  certain  that  it  is  not  the  same  as  P.  fallax  or  P.  duhius. 
It  does  somewhat  resemble  the  P.  neglectus  from  Trutta  fario  and  there 


180 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


[180 


is  a  possibility  that  it  may  be  the  same.  For  the  present  it  must  be 
considered  to  be  a  species  inquirenda.  In  regard  to  the  name  of  this 
species  attention  is  called  to  the  fact  that  Zschokke  wrote  the  name  as 
a  trinomial.  Since  a  trinomial  name  can  have  no  standing  and  since 
Monticelli  (1891)  first  wrote  this  name  as  a  binomial  it  is  here  sug- 
gested that  the  name  should  be  credited  to  him.  The  species  is  then  to 
be  known  as  Proteocephalus  salmonis-umblae  (Monticelli).  Pallas  (1811- 
12)  is  unaccessible  to  the  writer.  It  appears  that  his  description  of 
Taenia  salmonis  omul  if  one  were  given  was  of  no  value.  Monticelli 
(1891)  suggested  that  Taenia  salmonis-umhlae  was  probably  identical 
with  the  Taenia  salmonis-omul  Pallas  and  he  called  the  latter  a  species 
dubia.  Rudolphi  (1819:175)  expressed  the  opinion  that  Taenia  salmonis 
omul  was  a  species  of  Bothriocephalus.  For  this  reason  it  seems  that 
Taenia  salmonis  omul  is  hardly  of  sufficient  value  as  a  name  to  be 
worthy  of  being  called  a  species  inquirenda  and  since  Monticelli  ap- 
parently desired  to  bury  it  with  the  Taenia  salmonis-umhlae  it  is  best 
not  to  attempt  to  resurrect  it.  La  Rue  (1911:475)  in  a  list  of  Proteo- 
cephalus species  credited  this  species  to  Monticelli  who  was  the  first  to 
give  it  a  binary  name. 

PROTEOCEPHALUS  OSCULATUS  (Goeze),  sp.  inq. 
[Figs.  161-165,  182] 


1782: 

Taenia  osculata 

Goeze 

1782 :415 

1782: 

Taenia  alternatim  transverse  lineata 

Goeze 

1782 :416 

1786: 

Taenia  sUuri 

Batsch 

1786 :157-159 

1786 : 

Taenia  osculata 

Batsch 

1786 :209 

1788: 

Taenia  osculata 

Schrank 

1788 :47 

1788: 

Taenia  glanis 

Schrank 

1788 :47 

1790: 

Taenia  percae  p 

Gmelin 

1790:3079 

1790: 

Taenia  sUuri 

Gmelin 

1790:3080 

1803: 

Halysis  siluri 

Zeder 

1803 :353 

1803: 

Halysis  transverse-lineaia 

Zeder 

1803:353 

1810: 

Taenia  calycina 

Rudolphi 

1810:115-116 

1810: 

Taenia  osculata 

Rudolphi 

1810 :116-119 

1819: 

Taenia  osculata 

Rudolphi 

1819:150,497 

1845: 

Taenia  osculata 

Dujardin 

1845 :584 

1850: 

Taenia  osculata 

Diesing 

1850:522-523 

1854: 

Taenia  osculata 

Wagener 

1854 

1861: 

Taenia  osculata 

Van  Beneden  1861 :165 

1896: 

Ichthyotaenia  osculata 

Riggenbach 

1896 :267 

1909: 

Ichihyotaenia  osculata 

Liihe 

1909 :30-31 

1911: 

Proteocephalus  osculatus 

La  Bue 

1911 :475 

181] 


PROTEOCEPHALIDAE—LA  RUE 


181 


Specific  Diagnosis:    For  the  specific  diagnosis  see  Liihe's  diagnosis 
of  this  species  at  the  close  of  the  historical  summary. 

Habitat:    In  intestine  of  Silurus  glanis  L. 


Host 

Locality 

Magdeburg 
Greifswald 

Collector 

Authority 

Silurus  glanis  L. 
Silurus  glanis  L. 

Goeze 
Rudolphi 

Goeze         1782:415-416 
Rudolphi     1819:150 

Goeze  (1782:415)  described  and  figured  this  form  from  the  intes- 
tine of  Silurus  glanis.    His  diagnosis  is  here  given : 

"Der  gemijndete  Bandwurm.  Taenia  osculata.  Auch  vom  Graf  von  Borke 
in  den  Gedarmen  eines  24  pfiindigen  Welses  (Silurus  Glanis  L.).  Nach  der  Grosse 
dieses  Fisches  wenige,  und  ausserst  kleine  Tanien.  Kleiner,  als  in  der  Hechten. 
Nicht  am  Magenende,  sondern  mehr  in  der  Gegend  des  Afters.  Das  merkwurdigste 
daran,  das  sie  ausser  den  beyden  Saugblasen  am  Kopfe,  an  dem  vorstehendem 
Russel  eine  wahre  Miindung  batten,  welche  unter  dem  Komposito  mit  No.  4  Tub. 
B.  deutlich  zu  sehen  war.  An  keinem  andern  hat  der  Graf  die  Riisselmiindung  so 
deutlich,  als  an  diesen  wahrgenommen.  Sie  waren  alle  lebendig;  aber  auf  der 
Glasplatte  schwer  zu  behandeln,  weil  sie  sich  bestandig  kriimmten,  und  mit  ihren 
Saugwarzen  selbst  an  ihrem  eignem  Korper  vest  ansogen.  Ein  besondcre  Umstand ! 
Das  Aufblahen  der  Saugwarzen,  und  das  Ausstrecken  und  Einziehen  das  Riissels 
ein  angenehmes  Schauspiel.  Die  Glieder  sehr  undeutlich.  Mehr  Runzeln  als 
Glieder." 

Goeze  (1782:416)  described  a  form  from  Silurus  glanis  which  he 
thought  was  a  species  distinct  from  the  Taenia  osculata.  Later  investi- 
gators considered  that  the  second  form  was  identical  with  Taenia  oscu- 
lata and  it  seems  that  they  are  probably  correct.  Goeze 's  description  of 
the  second  form  is  more  complete  than  that  of  the  first  hence  it  is  here 
quoted : 

"Der  wechselsweise  Lineirte  Bandwurm.  Taenia,  alternatim  transverse  line- 
ata:  articulis  quadrangularibus ;  capite  quadriverrucoso. 

"Eine  wirklich  neue  und  besondere  Art.  Auch  aus  den  Gedarmen  eines 
sechzehnpfiindigen  Welses  (Silurus  Glanis.  L.)  Als  ich  am  isten  lulius  1780  von 
Rekane  nach  Magdeburg  kam,  blieb  ich  daselbst  um  der  grosseren  Fische  willen 
nach  einige  Tage.  Ich  bekam  die  Gedarme  eines  Stors  (Acipenser  Sturio  L.)  von 
50  Ffund.  In  denselben  verschiedene  Askariden,  und  zween  Kratzer,  wie  bey 
andern  Fischen.  In  den  Gedarmen  eines  Welsens  aber  fand  ich  zween  merk- 
wiirdige  Bandwurmer. 

"Die  Lange  etwa  12  Zoll.  Die  Breite  am  breitestem  Hinterende  eine  gute 
Linie.  Die  Glieder  regelmassige  Quadrate:  eine  Linie  lang  und  Breit.  Nach  dem 
Kopfe  zu  Verhaltnissmassig  kleiner.     Das   Karakteristische  dieses  Wurms  waren 


,  -I  <S 


182  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [182 

die,  Wechselsweise,  in  jedem  Gliede,  in  ungleicher  Ordnung  stehende  weisse  Quer- 
linien,  die  bis  iiber  die  Halfte  des  Korpers,  vom  Hinterende  an,  hinaufgehen,  und 
besonders  in  die  Augen  fallen.  Diese  Linien  gehen  nicht  weiter  bis  in  die  Mitte 
des  Gliedes.  Zeigt  sich  z.  B.  auf  der  rechten  Seite  in  jedem  Gliede,  in  fiinfen  nach 
einander,  die  weisse  Querlinie ;  so  ist  die  linke  Seite  f rey.  Dann  auf  der  linken 
Seite  in  zwey  Gliedern,  und  die  rechte  Seite  frey.  Weiter  nur  an  einem  Gliede, 
bald  rechts,  bald  links,  u.  s.  w.  In  der  Mitte  jedes  Gliedes  am  Hinterende  liegen 
reife  Eyer,  und  ich  kann  diese  Linien  fiir  nichts  anders,  als  fiir  Eyergange  halten, 
obgleich  die  Randmiindungen  zu  klein,  oder  zu  \erschlossen  sind,  als  dass  man  sie 
wahrnehmen  konnte.  Das  Kopfende  sehr  fein  und  diinne,  aber  flach,  und  das 
Kopfchen  nach  Proportion  wie  ein  kleines  Knotgen.  Unter  dem  Komposito 
der  Hals  ungegliedert.  Am  Kopfe  auf  jeder  Seite  zwo  stark  hervorstehende  Saug- 
miindungen.     Vom  ein  kurzes  Russelchen.  ohne  Baken." 

The  "  Querlinien "  of  the  above  description  is  probably  the  more  or 
less  clear  region  in  which  the  cirrus-pouch,  vagina,  and  the  convoluted 
vas  deferens  lie. 

Goeze's  figures  of  this  form  have  been  reproduced  (Figs.  163,  164, 
165). 

Batsch  (1786)  followed  Goeze  in  considering  that  two  species  of 
Taenia  were  parasitic  in  SUurus  glanis.  He  changed  the  name  of 
Goeze's  second  form  to  Taenia  siluri.  His  two  descriptions  which  seem 
to  be  based  largely  on  the  work  of  Goeze  are  here  quoted : 

"Taenia  siluri.    Der  Welsbandwurm. 

"Goetze,  S.  416.  T.  33.  fig.  11-14.  Taenia  (scalaris)  ductu  rectissimo  brevi 
introrsum  subclavato,  extrorsum  pedicellato ;  capite  papillis  quatuor  et  nodo  verti- 
cali  subaequalibus  globosis ;  artieulis  subquadratis,  marginibus  convexis. 

"In  jedem  reifen  Gliede  hat  der  Eiergang,  fast  wie  bei  dem  vorigen,  die 
Gestalt  eines  Stempels,  in  dem  er  mit  dem  Eierstocke,  welcher  in  der  Mitte  des 
Gliedes  liegt  und  eiformig  ist,  zusammenfliesst.  Aber  der  Gang  ist  kurzer,  gerader, 
und  der  Eierstock  weder  hervorragend  noch  so  ausgebreitet  und  deutlich  von  dem 
Eiergange  an  Grosse  verschieden. 

"Die  Glieder  sind  viereckig,  eine  Linie  lang  und  breit,  und  an  den  Ecken 
abgerundet,  und  an  den  Seiten  mit  einem  ziemlich  breiten  dunkeln  Rande. 

"Die  Eierstocke  und  Gange  sind  von  weisser  Farbe,  und  stehen  wechselsweise 
auf  einer  oder  der  andern  Seite,  in  verschiedner  Anzahl  neben  einander,  wie  bei 
den  vorigen  Art.  Sie  sind  vom  Hinterende  bis  iiber  die  Halfte  des  Korpers  sicht- 
bar,  wo  die  Glieder  aufhoren  viereckig  zu  sein,  sondern  ganz  kurz  werden,  und 
auch  von  da  an  mit  dem  ganzen  Vorderende  des  Korpers  gleichformig  bis  an  den 
Kopf  an  Breite  abnehmen.  Nahe  am  Kopfe  hat  der  Korper  etwa  den  sechsten 
Theil  der  Breite  der  reifen  Hinterhalfte.    Das  Vorderende  ist  flach. 

"Der  Hals  ist  lang  und  ungegliedert;  der  Kopf  ragt  wie  ein  starker  Knoten 
hervor,  der  aus  vier  kuglichen  grossen  nahe  beisammen  stehenden  Saugblasen,  und 
einem  beinahe  eben  so  grossen  Knopfchen  auf  dem  Scheitel  zusammengefesst  ist. 


183]  PROTEOCEPHALIDAE—LA  RUE  183 

"Jede  Saugblase  hat  einem  breiten  wulstigen  Rand  um  ihre  tiefe  Hohle;  und 
giebt  dem  Kopfe  eine  ausgezeichnete  Schonheit. 

"Die  Lange  des  ganzen  Korpers  erstreckt  sich  auf  zwolf  Zoll.  Diese  Art 
wohnt  in  den  Gedarmen  des  Welses  (Silurus  Glanis),  Goetz  hat  zwei  Zeichnungen 
davon  gegeben,  deren  eine  darin  von  dem  beschriebnen  abweicht,  dass  die  Hinter- 
glieder  zwar  viereckig  sind,  aber  doch  mit  dem  etwas  breitern  Hinterrande  iiber 
den  folgenden  Vorderrand  eckig  hervorstehen,  und  keine  Eiergange  zeigen.  Der 
Kopf  ist  iibrigens  in  der  Hauptsache  nicht  verschieden. 

"Die  Oeffnungen  der  Eiergange  und  die  Art  der  Eier  selbst  ist  an  dieser  Art 
noch  nicht  berichtigt." 

"Taenia  osculata.    Der  gentundete  Bandwurm. 

"Gotze  S.  41 5,  T.  Z3-  fig-  9-  10. 
Taenia   (larvata)   capite  cum  coUo  coalito,  osculis  binis  speciosis  bimarginatis,  m 
acumen  osculatum  defluente ;  corpore  crenato,  articulis  brevibus,  parallelepipedis. 

"Dieser  Bandwurm,  den  der  Graf  Borke  im  Wels  endeckt  hat,  ist  von  einer 
sehr  sonderbaren  Gestalt.  Er  hat  an  seinem  Kopfe,  der  fast  nicht  vom  Korper 
unterschieden  ist,  zwei  (vermuthlich  vier)  sehr  grosse  rundliche  Saugblasen,  welche 
dicht  an  einander  liegen,  die  ganze  Breite  des  Kopf  erfiillen,  und  mit  doppelten 
wulstigen  Randern,  wie  mit  Augensternen  versehen  sind.  Der  aussere  Wulst  ist 
nach  der  Zeichnung  dunkler  als  der  innere,  und  gefleckt,  beide  aber  etwas  gestrahlt. 
Mitten  zwischen  ihnen  kann  sich  der  Scheitel  in  eine  zapfen-  oder  schlauchahnliche 
kegelartige  Erhohung  verlangern,  welche  am  Ende  eine  Miindung  hat,  unten  sicb 
in  den  Kopf  ausbreiter,  und  mit  ihm  zusammenfliesst  Man  diirfte  auf  den  Ge- 
danken  gerathen,  als  wenn  dieser  Wurm  von  dem  oben-beschriebnen  Welsband- 
wurme  nicht  wesentlich  verschieden  ware,  allein  man  hat  bei  jenem  die  kegel- 
f ormige  Verlangerung  des  Scheitels  und  seine  Miindung  nicht  bemerkt ;  auch  sind 
bei  eben  demselben  die  Glieder  grosser  im  Verhaltniss  des  Korpers  und  deutlich 
organisirt.  Hier  ist  diess  nicht,  sie  gleichen  mehr  undeutlichen  Runzeln,  und  nach 
der  Vergrosserung  sind  sie  viereckig,  fiinfmal  breiter  als  lang,  mit  abgestumpften 
Erken.     Der  Hals  ist  ungegliedert  und  punktirt. 

"Es  waren  wenige,  ausserst  kleine  Bandwiirmer  im  Verhaltniss  gegen  den 
grossen  Fisch,  kleiner  als  im  Hecht.  Sie  befanden  sich  mehr  in  der  Gegend  des 
Afters,  kriimmten  sich  im  Leben  bestandig,  und  sogen  sich  mit  den  Saugwarzen 
an  ihrem  eigenen  Korper  an.  Das  Ausstrecken  und  Einziehen  des  Riissels  wie 
auch  das  Aufblahen  der  Saugwarzen,  gab  ein  angenehmes  Schauspiel." 

Schraiik  (1788)  devoted  a  two  line  diagnosis  to  each  of  the  two 
forms  reported  by  Goeze  from  Silurus  glanis.  The  second  form  he 
called  Taenia  glanis.  His  diagnosis  adds  nothing  to  the  data  given  by 
Goeze  and  Batsch.  Gmelin  (1790)  gave  Latin  diagnoses  for  Taenia 
siluri  and  Taenia  percae  form  p.  These  diagnoses  are  of  no  value  except 
to  show  that  he  regarded  these  two  forms  reported  by  Goeze  to  be  two 
distinct  species. 

Rudolphi  (1810:116-119)  summed  up  the  knowledge  of  this  species 
and  for  the  first  time  called  attention  to  the  fact  that  Goeze 's  two  forms 


184  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [184 

constituted  but  a  single  species.  Rudolphi  apparently  had  a  personal 
knowledge  of  this  species  and  hence  his  data  cany  much  more  weight 
than  do  those  of  some  of  the  other  writers  immediately  preceding  him. 
His  description  together  with  his  synonymy  is  here  quoted : 

"Taenia  osculata  Goeze. 

"Taenia:  osculis  rostellique  apice  concavis,  parte  antica  capillari,  articulif 
quadratis  plants,  margine  majorum  integerrimo. 

Goeze  Naturg.  p.  415.  Tab.  33.  fig.  9.  10.    T.  osculata. 

Idem  ib.  p.  416.  Tab.  33.  fig.  11-14.    Taenia  alternatim  transverse  lineata. 

Batsch  Bandw.  p.  209.  n.  45.  fig.  146.  147.  T.  osculata.  p.  157.  n.  16.  fig.  80. 
82.    T.  siluri. 

Schrank  Verzeichn.  p.  47.  n.  141.    T.  osculata.  n.  142.    T.  glanis. 

Gmel.  Syst.  Nat.  p.  3080.  n.  82.    T.  siluri.  ib.  p.  3079,  n.  77.    T.  percae  /3. 

Tabl.  Encycl.  t.  49.  fig.  4.  5.  (ic.  Goez.)  T.  siluri.  fig.  6-9.  (ic.  Goez.)  T. 
alternans. 

Zeder  Naturg.  p.  353.  n.  40.    Halysis  siluri.  n.  41.    Hal.  transverse-lineata, 

"Hab.  in  Siluri  Glanidis  parte  intestini  praesertim  media,  saepe  tamen  etiam 
inter  priorem.    Goezius  lulio,  ego  Majo,  reperimus. 

"Descr.  Vermes  pauci  a  Comite  de  Borke  reperti  exigui,  duo  a  Goezio  obser- 
vati  duodecim  pollices  longfi ;  mei,  copiosissimi,  pollicem  ad  pedem  et  quod  excurrit 
longi,  antice  capillares,  postice  lineam  ad  sesquilineam  lati. 

"Caput  discretum,  subglobosum,  exiguum,  Oscula,  uti  caput  totum,  quam  in 
specie  praecedente  multo  minora,  concava,  profunda,  ostio  exiguo,  ut  in  ilia  sita 
saepeque  simul  in  conspectum  venientia.  Rostellum  intermedium,  brevissimum, 
osculum  quintum,  sed  minus  et  plerumque  magis  protrusum  refert.  Corporis  plani 
et  tenuis  pars  antica  in  verme  tam  poUicari  quam  pedali  capillaris,  mox  tantum 
rugosa  vel  incisa  videtur  colli  speciem  sistens,  mox  autem  articulis  distinctis  iisque 
tenuissimis,  forma  variis,  m<iniliformibus,  oblongis  et  rugaeformibus  interjectis, 
constat ;  pars  reliqua  sensim  increscens,  articulis  tandem  quadratis,  satis  aequalibus, 
margine  laterali  rectiusculo,  integerrimo.  Articulos  ultimus  rotundatus,  fere  sem- 
per emarginatus.  Cujusvis  articuli  majoris  media  pars  ovariutn,  sive  maculam  ob- 
longam,  pellucidam  continet,  quae  neque  anteriorem,  neque  posteriorem  articuli 
marginem  attingit.  A  quolibet  ovario  linea  transversa  ad  alterum  articuli  margi- 
nem  lateralem,  ordine  plerumque  alterno,  non  tamen  certo,  excurrit.  Linea  longi- 
tudinalis,  qualis  in  specie  praecedente,  in  hac  non  conspicua. 

"Substantia  vermis  tenuis,  at  firma  et  duriuscula,  ideoque  in  aqua  non  pro 
speciei  praecedentis  more  intumescit  vel  gelatinosa  fit.  Vermibus  utriusque  speciei 
aeque  magnis,  prioris  volumen  multo  majus. 

"Obs.  I.  Hujus  et  praecedentis  discrimen,  quo  minus  conjungantur,  nimis 
magnum  esse,  cuique  observationem  illi  adjectam  legenti,  vel  utramque  recentem  aut 
spiritu  vini  servatum  comparanti  patebit. 

"Obs.  2.  Me  speciem  a  Comite  de  Borke  detectam  et  Goezio  osculatam  dic- 
tam,  cum  altera  hujus  auctoris  specie  (transversim  lineata)  conjunxisse  forsan 
miveris,  sed  utrique  collum  non  articulatum  et  tenue  adscribitur,  quale  praecedenti 


185]  PROTEOCEPHALIDAE—LA  RUE  185 

semper  deest,  et  Borkius,  qui  Taeniolas  suas  in  Glanidis  intestinis  crassis  ^epererit, 
nonnisi  partem  vermium  summam  offendisse  videtur,  dum  articulos  rugas  potius 
referre  asserit,  quod  de  hujus  tantum  parte  antica  valet.  Goezius  vermes  majores 
reperit  et  ejusdem  icon  (fig.  ii.).  Taeniam  integram  sistens,  bona  est.  Capitis 
vero  icones  neque  Borkiana  neque  Goeziana  laude  dignae  sunt.  Zederus,  verme 
licet  non  vise,  alteram  Goezii  speciem  rescindendam  esse,  recte  praedixerat. 

"Obs.  3.  Miillerus  de  Taenia  ocellata,  nobis  n.  21.  dicta,  loquendo,  Taeniam 
transverse-lineatam  Goezii  ab  ilia  lineolarum  lateralium  defectu  solo  forsan  dif- 
ferre,  suspicatus  est,  quo  Gmelinus  commotus,  qui  Taeniam  nostram  oscellatae 
varietatem  fingeret,  sed  osculata  toto  coelo  diversa,  et  rostello  solo  jam  distingui 
potest. 

"Obs.  4.  Omnes  quidem  Taeniae  osculatae  sunt,  nostra  autem  (uti  et  praece- 
dens)  Kar'i^oxrjO  ita  \T)cari  potest,  cum  osculo  quinto  quasi  instruatur,  et  oscula 
valde  profunda  sint.  Goezii  ergo  nomen  conservavi,  et  praecedentem  simili  voce 
designavi." 

Rudolphi  (1810:115)  also  described  Taenia  calycina  from  Silurus 
glanis.  This  species  he  later  (1819:497)  came  to  consider  as  identical 
with  Taenia  osculata.    His  later  observations  are  here  given : 

"Entozoologiam  edendo  duplicem  hujus  Taenia  formam,  mihi  tum  temporis 
constantem  visam  prae  oculis  habui,  quo  commotus  T.  calycinam  mihi  dictam  ab 
osculata  distinxi.  Postmodum  vero  Gryphiae  Octobri  mense  Siluri  Glanidis  quin- 
quaginta  tres  libras  pondere  aequantis  intestina  examinando,  Taenias  plurimas 
reperi  inter  T.  osculatam  et  calycinam  et  ita  quidem  intermedias,  ut  nullum  amplius 
discrimen  superesset.  Tres  quatuorve  pedes  longae  collo  mox  longiore  mox  bre- 
viore  utebantur;  articuli  lineis  longitudinalibus,  rarius  (posteriores)  etiam  trans- 
versis  insignes ;  aquae  commissae  quasi  gelatinosae  reddebantur,  aliquot  dies  vivae 
conservatae.    Taenia  calycina  ideoque  rescindenda  est." 

Rudolphi  (1819:150)  gave  a  very  brief  Latin  diagnosis  of  T.  oscu- 
lata and  stated  that  he  had  collected  it  at  Greifswald.  Dujardin  (1845) 
and  Diesing  (1850)  added  very  little  to  the  previous  diagnoses  and  de- 
scriptions. Wagener  (1854)  is  inaccessible  to  the  writer  hence  it  is 
impossible  to  review  his  findings.  Cams  (1857)  gave  two  drawings  of 
this  form  after  Wagener  (1854).  Van  Beneden  (1861:165)  listed  this 
species  of  cestode.  Riggenbach  (1896)  considered  it  a  species  of  Ich- 
thyotaenia  but  he  gave  no  descriptive  data  on  the  species.  Two  draw- 
ings of  the  head  by  Wagener  are  reproduced  (Figs,  161,  162). 

Liihe  (1909:30-31)  gave  a  short  descriptive  diagnosis  of  the  species 
and  in  an  original  figure,  reproduced  (Fig.  182),  he  showed  a  little  of 
the  structure  of  the  proglottid.    His  diagnosis  reads: 

"Zirka  50  cm  bis  i  m  lang,  bei  einer  grossten  Breite  von  ca.  3  mm.  Scolex 
mit  einem  kleinen  scheitelstandigen  "Saugnapf",  der  mit  zahlreichen,  sehr  hinfalli- 
gen   Hakchen   von  0,007   niin   Lange   besetzt   ist.     Proglottiden    mit   entwickelten 


186  ILUNOIS  BIOLOGICAL  MONOGRAPHS  [186 

Geschlechtsdrusen  sowie  auch  solche  mit  Uterus  annahemd  quadratisch.  Genital- 
offnung  in  der  Mitte  des  Gliedrandes.  Cirrusbeutel  klein,  kaum  uber  den  Dot- 
terstock  nach  innen  hinausragend ;  das  Knauel  des  Vas  deferens  stark  querge- 
streckt,  vom  Cirrusbeutel  bis  an  die  Medianlinie  reichend.  Hodenblaschen  sehr 
zahlreich,  das  ganze  Mittelfeld  der  Proglottis  einnehmend.  Vagina  ohne  Sphincter. 
Uterus  jederseits  mit  etwa  6-8  schlanken,  sich  etwas  verzweigenden  Seitenasten. 
Kalkkorperchen  anscheinend  sparlich. 
"Im  Darm  von  Silurus  giants  L." 

La  Rue  (1911:475)  listed  this  as  a  little  known  species  of  Proteo- 
cephalus. 

The  above  diagnosis  by  Liihe  and  his  drawing  of  the  inner  struc- 
ture of  the  proglottid  furnish  the  only  available  data  on  the  anatomy 
of  this  species.  Unfortunately  his  drawing  and  description  fail  to  show 
many  of  the  details  which  one  might  desire.  If  his  interpretation  of  the 
structures  there  found  are  correct  then  one  must  consider  that  so  far 
as  the  arrangement  and  character  of  the  generative  organs  are  con- 
cerned this  species  must  belong  with  the  genus  Proteocephalus.  The 
presence  of  the  minute  spines  on  the  fifth  sucker  is  not  in  strict  accord 
with  the  writer's  ideas  of  this  genus.  At  first  glimpse  one  is  caused  to 
think  of  the  spines  on  the  head  of  the  members  of  the  genus  Acantho- 
taenia  and  is  led  to  inquire  whether  this  species  does  not  have  its  place 
in  that  genus.  However,  since  the  Acanthotaenia  are  found  thus  far 
only  in  the  Varanidae,  a  family  of  lizards,  it  seems  that  this  species 
can  not  have  its  closest  relatives  there.  Since  its  host  is  a  member  of 
the  genus  Silurus  and  since  this  genus  harbors  several  species  of  ces- 
todes  of  the  genus  Monticellia  one  must  consider  the  possibility  of  its 
belonging  with  the  latter.  Here,  it  seems  to  the  writer,  this  species  will 
eventually  be  placed  but  at  present  aU  the  data,  meager  tho  they  are  in 
details,  place  it  in  the  genus  Proteocephalus. 

TAENIA   SIMPLICISSIMA  Leidy 

1887 :  Taenia  simplicissima  Leidy  1887 :22 

1891:  Taenia  simplicissima  MonticeUi  1891 

1896 :  Ickihyoiaenia  simplicissima       Riggenbach  1896 :267 

1911:  Proteocephalus  simplicissimtis  La  Rue  1911:475 

This  species  was  first  described  in  a  very  superficial  way  by  Leidy 
(1887)  who  besides  the  wholly  inadequate  species  description  figured 
the  head  and  the  outline  of  the  proglottids  in  two  drawings.  Monticelli 
(1891)  without  comment  included  this  species  in  a  list  of  forms  which 
somewhat  resembled  his  Teiracotylus  (Taenia)  coryphicephala.  Rig- 
genbach (1896)  included  this  form  in  a  list  of  species  of  Ichthyotaenia. 


187]  PROTEOCEPHALIDAE—LA  RUE  187 

He  did  not  discuss  his  reasons  for  so  doing.     La  Rue   (1911)   inad- 
vertently included  this  form  among  a  list  of  species  of  Proteocephalus. 
Leidy's  description  is  here  quoted: 

"Taenia  simplicissima. — Head  small,  unarmed,  truncate;  bothria  spherical,  ter- 
minal, occupying  the  four  angles;  neck  very  long,  nearly  or  as  wide  as  the  head, 
body  gradually  widening  to  the  posterior  third  and  then  tapering;  anterior  seg- 
ments transversely  linear,  subsequently  reversed  disklike,  gradually  longer  and 
wider,  then  campanulate  and  gradually  becoming  longer  and  narrower.  Generative 
apertures  and  ova  unobserved.  A  number  of  specimens  from  the  Cod,  Gadus 
callarias,  up  to  20  lines  by  i  mm.  where  widest.  Two  only  of  the  specimens  re- 
tained the  head." 

The  only  reliable  data  in  the  description  and  the  drawings  have  to 
do  with  the  head  and  the  form  of  the  proglottids.  Not  even  the  genital 
pores  were  noted.  Furthermore  so  far  as  the  writer  is  able  to  find  out 
no  investigator  since  Leidy  has  seen  or  described  this  parasite  of  the 
cod.  Nor  is  it  likely  that  Leidy's  specimens  remain  for  he  did  not  save 
many  of  them  and  many  of  those  which  he  preserved  have  since  been 
destroyed  hence  a  re-examination  of  his  specimens  seems  unlikely.  Since 
the  host  is  a  purely  marine  fish  and  since  no  undoubted  species  of 
Proteocephalus  are  known  to  come  from  marine  hosts  the  writer  holds 
it  highly  improbable  that  this  form  is  a  species  of  Proteocephalus. 
Other  than  the  fact  that  it  has  four  sessile  suckers  and  inhabits  a  fish 
it  certainly  has  no  claim  for  a  place  in  the  genus  Proteocephalus.  The 
species  is  therefore  deleted  from  the  list  of  Proteocephalus  species  and 
Leidy's  original  name  for  it  is  retained. 

TAENIA  BELONES    Miiller 

The  writer  has  examined  the  statements  of  Rudolphi  (1819:175)  in 
regard  to  Taenia  belones  and  he  agrees  with  Monticelli  (1891)  and  Rig- 
genbach  (1896)  in  believing  that  this  form  does  not  properly  belong  in 
the  genus  Proteocephalus.  Monticelli  (1891)  expressed  the  view  that 
this  species  is  identical  with  Bothriocephalus  ielones  Dujardin.  Rig- 
genbach  (1896)  apparently  accepted  Monticelli 's  statement. 

TAENIA  POLLACHII  Rathke 

Rudolphi  (1819:175)  gave  a  short  statement  in  regard  to  this  form 
but  evidently  he  did  not  regard  it  as  a  well  known  species.  Monticelli 
(1891)  thought  it  very  probable  that  this  form  was  identical  with 
Abothrium  gadi  Van  Beneden.  Riggenbach  (1896)  apparently  ac- 
cepted this  view.  The  writer  has  found  no  reason  for  thinking  that  this 
form  is  a  species  of  Proteocephalus. 


188  ILUNOIS  BIOLOGICAL  MONOGRAPHS  [188 

CHOANOSCOLEX  ABSCISUS  (Riggenbach  1896)  La  Rue 

[Fig.  151]    , 

1896:    Ichthyotaenia  ahscisa  Riggenbach        1896:193-210 

1911:     Choanoscolex  ahscisus  La  Rue  1911:479 

Specific  Diagnosis :  Characters  of  genus.  Worms  small,  25-30  mm. 
long,  breadth  up  to  0.765  mm.  Number  of  proglottids  as  many  as  70. 
Head  somewhat  conical  with  a  fold  of  tissue  at  the  base  at  times  partly 
covering  the  suckers.  Apex  of  head  flattened.  Maximum  breadth  of 
head  0.476  mm.  No  rostellum,  no  fifth  sucker.  Suckers  large  oval, 
0.306  mm.  long  by  0.136  mm.  broad.  First  proglottid  longer  than  broad. 
Tenth  proglottid  0.476  mm.  broad  by  0.102  mm.  long.  Segments  near 
middle  of  worm  broader  than  long  or  nearly  quadrate.  Last  proglottid 
1.02  mm.  long  by  0.765  mm.  broad.  Proglottid  margins  straight,  pos- 
terior comers  rounded.    Last  ploglottid  with  bluntly  rounded  end. 

Sexual  organs  as  in  genus.  Genital  aperture  situated  at  end  of 
anterior  Vq-Yq  of  the  proglottid.  No  genital  papilla.  Testes,  about  100 
in  number,  0.054  by  0.045  mm.  in  size,  arranged  in  a  single  layer  be- 
tween vitellaria.  Coils  of  vas  deferens  voluminous.  Coils  of  ductus 
ejaculatorius  few.  Cirrus  swollen,  with  large  lumen.  Cirrus-pouch 
large,  pear-shaped,  muscular,  reaching  nearly  to  middle  of  segment. 
Length  of  cirrus-pouch  0.34  mm.  Vagina  anterior  or  posterior  to  cirrus- 
pouch.  Lumen  of  first  part  of  vagina  large.  Receptaculum  seminis 
present.  Ovary  bilobed,  posterior,  unbranched.  Vitellaria  lateral,  fol- 
licular. Uterus  a  median  tube  with  many  lateral  outpocketings.  Eggs, 
0.021  mm.  long  by  0.016  mm.  broad. 

Habitat:    Silurus  sp.,  Rio  Paraguay,  South  America. 

This  species  was  described  by  Riggenbach  (1896)  as  a  species  of 
Ichthyotaenia.  La  Rue  (1911:479)  established  for  it  a  new  genus, 
Choanoscolex  and  made  this  species  the  type  of  the  new  genus. 

The  material  was  collected  by  Dr.  Ternetz  in  Paraguay,  1894.  The 
following  description  is  based  on  the  work  of  Riggenbach  (1896).  Rig- 
genbach considered  this  form  to  be  a  species  of  Ichthyotaenia  and  his 
comparisons  are  made  with  species  of  that  genus  rather  than  with 
C  orallobothrium . 

The  worm  is  small  being  25-30  mm.  in  length.  It  is  made  up  of 
about  70  proglottids  whose  form  varies  according  to  the  age.  The  scolex 
(Fig.  151)  varies  from  the  usual  shape  in  being  somewhat  conical  in- 
stead of  spherical  and  in  having  at  the  base  of  the  cone  a  fold  of  tissue 
or  a  mantle  which  may  be  partially  drawn  over  the  suckers.  The  tip 
of  the  cone  is  flattened.    The  maximum  breadth  of  the  head  is  0.476  mm.. 


189]  PROTEOCEPHALIDAE—LA  RUE  189 

the  breadth  of  the  flattened  apex  of  the  cone  0.255  mm.  The  suckers 
which  are  separated  from  each  other  by  a  small  zone  only  are  very  large 
in  proportion  to  the  size  of  the  head.  The  shape  is  oval  or  at  times 
triangular.  Their  length  is  about  0.306  mm.  and  their  breadth  about 
0.136  mm. 

The  head  reaches  its  greatest  breadth,  0.476  mm.,  at  the  point  where 
it  passes  over  into  the  neck  of  medium  length  and  a  breadth  of  0.348  mm. 
First  proglottids  are  very  narrow  transverse  bands.  The  tenth  pro- 
glottid is  about  0.476  mm.  broad  by  0.102  mm.  long.  As  in  most  species 
the  length  of  the  proglottid  increases  with  age  more  rapidly  than  the 
breadth.  The  twentieth  proglottid  has  a  length  of  0.153  mm.  Proglot- 
tids near  the  middle  of  the  worm  are  broader  than  long  or  quadrate. 
The  last  proglottid  is  0.765  mm.  broad  by  1.02  mm.  long.  Margins  of 
the  proglottids  are  straight  and  the  posterior  comers  of  the  proglottids 
are  rounded.  Except  for  the  slight  elevation  in  the  region  of  the  genital 
opening,  the  geometrical  form  of  the  proglottid  is  not  disturbed.  The 
last  proglottid  is  rounded  off  at  the  posterior  end. 

Riggenbach  (1896:196-199)  described  the  cuticula,  parenchyma, 
musculature,  and  nervous  system.  These  structures  are  typical  for 
Proteocephalids.  He  did  not  describe  the  musculature  of  the  head.  In 
the  head  the  small  space  between  the  suckers  is  almost  completely  filled 
with  vessels  of  the  excretory  system.  The  four  main  vessels  are  bent 
inward  as  they  enter  the  head.  A  circular  anastomosis  behind  the  suck- 
ers could  not  be  demonstrated.  In  the  short  apical  region  of  the  head 
the  vessels  are  never  capillaries  and  an  apical  capillary  plexus  such  as 
is  found  in  Corallobothrium  lohosum  does  not  occur.  Riggenbach  was 
not  able  to  find  a  direct  connection  of  the  vessels  of  the  head  or  of  the 
plexus  with  the  exterior.  Posterior  to  the  suckers  the  main  excretory 
vessels  pass  to  the  lateral  fields  of  the  neck  in  a  course  that  is  perpen- 
dicular to  the  longitudinal  axis  of  the  body,  then  they  bend  posteriad, 
decrease  in  size  and  extend  throughout  the  strobila  as  nearly  straight 
narrow  tubes.  In  the  region  of  the  cirrus-pouch  and  vagina  the  dorsal 
vessel  passes  above  and  the  ventral  vessel  below  these  organs.  The  ex- 
cretory vessels  lie  mesad  of  the  vitellaria  and  the  lateral  nerve  trunks. 
At  the  posterior  part  of  each  segment  the  main  excretory  vessels  are 
connected  by  a  transverse  anastomosis.  In  the  region  of  the  transverse 
anastomosis  are  canals  which  connect  the  main  vessels  with  the  exterior. 
These  canals  arise  directly  from  the  main  vessels  and  not  from  a  capil- 
lary network.  The  point  of  opening  to  the  exterior  is  on  the  surface 
near  the  posterior  angle  of  the  proglottid.  In  the  posterior  part  of  the 
end-proglottid  the  four  main  vessels  join  together  to  form  a  small  blad- 


190  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [190 

der  or  reservoir.  This  reservoir  opens  in  the  middle  of  the  rounded 
posterior  margin  of  the  end-proglottid. 

The  genital  aperture  is  marginal,  irregularly  alternating,  situated 
at  the  end  of  the  anterior  l^  to  y^  of  the  proglottid.  A  genital  papilla 
is  not  present  tho  the  margin  of  the  genital  sinus  is  slightly  swollen. 
In  structure  and  arrangement  the  sexual  organs  agree  thoroly  with  the 
general  relations  of  Proteocephalus.  Riggenbach  mentions  a  receptacu- 
lum  seminis  as  being  new  to  Ichthyotaenia  but  it  has  been  found  in 
several  species  of  Proteocephalus. 

The  testes  number  about  100.  They  measure  0.054  by  0.045  mm. 
They  lie  in  a  single  layer  in  the  medullary  parenchyma,  between  the 
viteUaria  and  anterior  to  the  ovaries.  The  vas  deferens  forms  a  volumi- 
nous knot  of  coils  outside  the  cirrus-pouch.  The  ductus  ejaculatorius 
forms  a  few  coils  in  the  cirrus-pouch  and  then,  undergoing  a  change  in 
histological  structure,  it  passes  over  into  the  cirrus,  the  basal  part  of 
which  is  much  broadened  to  form  a  roomy  ovoidal  vesicle.  The  cirrus 
somewhat  resembles  the  cirrus  of  Monticellia  malopteruri  (Fritsch). 
Riggenbach  did  not  see  the  protruded  cirrus.  The  cirrus-pouch  is  a 
large  pear-shaped  and  muscular  sac  extending  into  the  segment  perpen- 
dicular to  the  margin.  Its  length  is  about  0.34  mm.  Since  the  ripe 
proglottids  measure  about  0.70-1.0  mm.  broad  the  cirrus-pouch  must 
reach  from  l^  to  %  across  the  proglottid. 

The  vagina  opens  into  the  common  genital  sinus  anterior  or  poste- 
rior to  the  cirrus-pouch.  In  old  proglottids  the  beginning  part  of  the 
vagina  is  swollen  into  a  sac  nearly  as  large  as  the  cirrus-pouch  but  in 
young  proglottids  the  diameter  of  the  vagina  is  uniform.  In  its  course 
to  the  middle  of  the  proglottid  it  describes  an  arc,  then  it  bends  sharply 
and  passes  to  the  interovarial  space  in  a  spiral  or  sinuous  course.  The 
convolutions  of  vas  deferens  are  crossed  by  the  vagina  only  when  the 
latter  opens  anterior  to  the  cirrus-pouch.  A  receptaculum  seminis  is 
present  near  the  mid-piece  of  the  ovary.  This  is  more  than  a  broaden- 
ing of  the  vagina  at  this  point  for  in  Riggenbach 's  drawing  there  is 
shown  a  change  in  the  histological  structure  of  the  vaginal  wall.  The 
relations  of  the  organs  with  the  interovarial  space  is  similar  to  that  in 
species  of  Proteocephalus  and  need  not  be  discussed  here.  The  bilobed 
ovary  is  in  the  posterior  region  of  the  proglottid.  The  lobes  are  un- 
branched  saclike  structures  united  by  a  mid-piece  from  which  the  ovi- 
duct arises.  The  vitelline  glands  are  lateral  follicular  structures  ex- 
tending the  fuU  length  of  the  segment.  They  occupy  a  broad  zone 
laterad  to  the  excretory  vessels  and  the  nerve  trunks.  The  uterus  is  a 
median  longitudinal  tube  with  a  large  number  of  lateral  outpocketings 
on  either  side.     The  uterine  eggs  are  0.021  mm.  long  by  0.016  mm. 


191]  PROTEOCEPHALIDAE—LA  RUE  191 

broad.  "The  elongated  shell  is  very  thin  and  surrounds  in  part  the 
yet  undifferentiated  egg-cell  with  the  yolk-cells,  in  part  the  cell-groups 
which  are  the  results  of  the  first  development  stages."  Evidently 
Riggenbach  saw  no  embryos. 

In  life  the  heads  of  typical  species  of  Proteocephalus  are  extremely 
variable  in  form  passing  with  considerable  rapidity  from  one  contraction 
state  to  another.  By  the  contraction  of  longitudinal  muscles  extending 
into  the  apex  of  the  head  the  suckers  may  be  drawn  down  out  of  sight 
within  the  inflated  neck  region  but  there  are  no  folds  of  tissue  at  the 
base  of  the  head  within  which  the  head  may  retreat.  In  the  species 
just  described  this  fold  of  tissue  within  which  the  suckers  are  partially 
withdrawn  seems  to  be  a  constant  feature  of  the  scolex.  Thus  the  head 
differs  from  heads  of  species  of  Proteocephalus.  At  first  one  notes  cer- 
tain similarities  with  the  scolices  of  species  of  Corallohothrium.  Fur- 
ther consideration  of  the  structure  of  the  two  types  of  heads  convinces 
one  that  they  are  not  alike.  The  head  of  a  Corallohothrium  is  greatly 
flattened  anteriorly  and  the  suckers  are  directed  anteriad.  At  the  mar- 
gins of  the  flattened  apical  region  are  numerous  folds  and  lappets  which 
form  a  corolla-like  sheath  about  the  suckers.  In  this  species  the  head 
is  conical  and  the  suckers  are  directed  outward.  In  Corallohothrium 
lohosum  Riggenbach  (1896)  found  a  small  muscle-cross  connecting  each 
dorsal  sucker  with  the  ventral  sucker  opposite  it.  Riggenbach  did  not 
see  such  a  structure  in  his  sections  of  the  heads  of  Ichthyotaenia  ah- 
scisa.  It  is  doubtful  if  it  occurs  in  any  other  genus  known  at  present 
to  belong  in  this  family. 

This  species  may  be  considered  as  forming  a  transitional  stage 
between  Proteocephalus  and  Corallobothrium.  This  view  is  supported 
by  the  appearance  of  heads  of  plerocercoids  of  an  unknown  species  of 
Corallobothrium  found  encysted  in  the  liver  of  Ameiurus  melas  and  A. 
nehulosus  from  the  Illinois  river.  The  heads  of  these  plerocercoids  pos- 
sessed but  a  few  simple  folds  of  tissue  enveloping  a  part  of  the  head. 
When  alive  the  heads  were  somewhat  conical,  not  flattened  on  the  apex 
as  in  preserved  heads  of  the  adult  worms.  However,  a  marked  difference 
between  these  heads  and  the  heads  of  Choanoscolex  ahscisa  may  be  noted. 
In  the  plerocercoids  the  suckers  are  plainly  paired,  two  dorsal  and  two 
ventral,  while  in  this  species  the  head  is  but  slightly  flattened  dorso- 
ventrally  and  the  suckers  are  not  plainly  paired.  This  species  does  not 
belong  in  the  genus  Corallobothrium  or  Proteocephalus. 

The  foregoing  descriptions  of  species  of  Proteocephalus  are  briefly 
summarized  in  and  supplemented  by  the  following  comparative  table. 


192 


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204  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [204 

OPHIOTAENIA  PERSPICUA  La  Rue 
[Figs.  10,  102] 
1911 :    Ophiotaenia  perspicua        La  Rue        1911 :480 

Specific  Diagnosis:  Characters  of  the  genus.  Length  up  to  36.0 
cm.  Maximum  breadth  about  2.0  mm.  Head  conical,  flattened,  divided 
by  grooves  into  four  quadrants.  Breadth  of  head  0.355-0.408  ,nm., 
thickness  of  head  as  much  as  0.306  mm.,  length  about  0.270  mm.  Suck- 
ers four,  circular,  oval  or  at  times  somewhat  triangular,  0.105-0,170  mm. 
in  maximum  dimension.  Cavity  of  sucker  0.053-0.106  mm.  in  diameter. 
Neck  long,  5-7  mm.  in  length  by  0.170-0.425  mm.  broad.  First  proglot- 
tids  short.  Mature  proglottids  quadrate,  2.0  mm.  in  length  and  breadth, 
or  somewhat  longer  than  broad.  Ripe  proglottids  measure  as  much  as 
3.8  mm.  long  by  1.2  mm.  broad.  Segmentation  indistinct.  Proglottids 
attached  by  full  width.  Surface  smooth.  In  life  and  when  preserved 
in  formol  specimens  somewhat  translucent. 

Genital  pore  marginal,  irregularly  alternating,  situated  near  middle 
or  at  end  of  first  third  of  proglottid.  Vagina  anterior  or  posterior  to 
cirrus-pouch.  Testes  150-215,  oval  or  polygonal,  measuring  up  to  0.053- 
0.106  mm.,  situated  in  two  fields,  Vas  deferens  in  ripe  proglottids  a 
heavy  mass  of  coils  reaching  from  end  of  cirrus-pouch  to  the  mid-field. 
Cirrus-pouch  0.255-0.320  mm.  long  by  0.080-0.090  mm,  broad.  Ratio  of 
length  of  cirrus-pouch  to  proglottid  breadth  1 :4  to  1 :3.  Cirrus  slender, 
about  0.20  mm.  long  when  protruded.  Vagina  dUated  in  first  part,  not 
crossing  cirrus-pouch.  Sphincter  vaginae  and  receptaculum  seminis 
present.  Lobes  of  ovary  long,  flattened,  irregular  in  outline,  made  up 
of  anastomosing  tubules.  Oocapt  and  ootype  present,  Vitellaria  loose. 
Uterus  when  ripe  with  20-30  lateral  pouches  on  either  side.  Uterine 
pores  not  seen.  Eggs  provided  with  three  membranes,  outer  one  0.045- 
0.100  mm.  in  diameter.    Embryos  0.018-0,021  mm. 

Habitat:  Natrix  (Nerodia)  rhomhifer  Hallowell,  (type  host); 
Havana,  Illinois,  (type  locality). 

Type :  Unaccessioned  bottles  in  coll.  La  Rue,  also  bottle  in  Dr.  H. 
B.  "Ward's  collection.    Slides  of  same. 

The  host,  a  very  large  female,  was  caught  on  the  banks  of  the  Illi- 
nois river  at  Havana,  Illinois,  and  was  examined  for  parasites  July  9, 
1910.  From  the  intestine  57  cestodes  and  pieces  were  taken.  Of  these 
42  pieces  had  heads  attached.  No  strobilas  were  found  with  an  end- 
proglottid.  There  were,  however,  a  number  of  specimens  with  ripe  pro- 
glottids.   Because  of  their  translucent  appearance  in  life  and  when  pre- 


205]  PROTEOCEPHALIDAE—LA  RUE  205 

served  in  formol  the  writer  (1911)  has  proposed  the  specific  name 
Ophiotaenia  perspicua.  Other  specimens  of  this  species  were  furnished 
the  writer  by  Mr.  Herman  Douthitt  from  Natrix  (Nerodia)  rhombifer 
taken  in  Oklahoma,  June,  1910. 

La  Rue  (1911 :480)  described  this  species  in  a  preliminary  way  and 
made  it  the  type  of  his  new  genus  Ophiotaenia. 

Two  heads  of  the  lot  from  Oklahoma  measured  respectively  0.374 
mm.  and  0.340  mm.  broad  by  0.27  mm.  thick.  The  suckers  measured 
0.119  mm.  long  by  0.102  mm.  broad.  The  neck  was  0.272  mm.  broad 
and  several  millimeters  long.  The  longest  worm  was  135  mm.  long  by 
a  maximum  breadth  of  1.19  mm.  Specimens  were  mounted  in  toto  and 
from  them  a  positive  determination  was  made.  The  longest  specimen 
of  the  preserved  type  material  measured  36.0  cm.  in  length  by  a  maxi- 
mum breadth  of  2.0  mm.  The  neck  is  long,  slightly  broader  than  the 
head  and  not  easily  distinguished  from  the  segmented  portion  following. 
The  first  proglottids  are  very  short.  About  16.0  cm.  from  the  head  the 
proglottids  are  quadrate,  length  and  breadth  being  about  2.0  mm.  These 
proglottids  are  mature.  At  26.0  cm.  from  the  head  the  proglottids 
begin  to  increase  in  length  and  to  decrease  in  breadth.  The  second  from 
the  last  proglottid  measures  3.8  mm.  long  by  1.2  mm.  broad.  These 
proglottids  are  ripe.  An  examination  of  other  specimens  of  the  lot 
shows  a  considerable  variation  in  dimensions.  Proglottids  are  attached 
by  their  full  width.  The  margins  of  the  strobila  are  smooth,  almost 
without  indentations  at  the  junction  of  the  proglottids.  As  a  conse- 
quence the  segmentation  is  indistinct  so  that  without  staining  and  clear- 
ing the  proglottid  limits  are  made  out  with  difficulty  even  with  a 
microscope.  The  proglottids  are  rectangular  in  shape,  rarely  oval.  The 
surface  of  the  worm  is  sometimes  thrown  into  shallow  longitudinal  fur- 
rows. No  transverse  furrows  are  noted.  In  life  and  when  preserved  in 
formol  the  specimens  are  translucent  from  which  character  is  derived 
the  specific  name. 

The  head  is  somewhat  conical  in  shape,  slightly  flattened  dorso- 
ventrally  and  marked  off  into  four  quadrants  by  grooves  which  extend 
from  the  basal  region  to,  or  nearly  to,  the  apex.  Each  quadrant  bears 
at  its  thickest  and  broadest  region  a  sucker  which  opens  outward  and 
slightly  forward.  The  apex  does  not  exhibit  a  circular  depression  or  a 
fifth  sucker.  The  head  varies  in  breadth  from  0.255  mm.  to  0.408  mm. 
This  dimension  slightly  exceeds  the  thickness  and  length.  A  head  0.357 
broad  measured  0.306  mm.  in  thickness,  while  a  head  0.408  mm.  broad 
was  about  0.270  mm.  long. 

The  suckers  show  also  considerable  variation  in  size  and  shape.  In 
general  they  are  nearly  round  or  oval  in  outline  with  shallow  cavities. 


206  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [206 

However  some  of  them  appear  nearly  triangular.  They  measure  0.105- 
0.170  mm.  in  diameter.  In  the  smaller  ones  the  effect  of  contraction 
is  very  plainly  shown  by  the  deeper  cavity  and  by  the  thickened  muscu- 
lar wall.  The  diameter  of  the  sucker  opening  varies  from  0.053  to 
0.105  mm.  A  vestigial  fifth  sucker  resembling  that  of  0.  filaroides  is 
found  imbedded  in  the  tissues  of  the  head.  The  neck  is  always  long, 
5  to  7  mm.,  and  its  breadth  is  subject  to  considerable  variation,  from 
0.170  to  0.425  mm. 

Since  this  study  is  based  upon  toto  preparations  almost  exclusively, 
the  excretory,  nervous  and  muscular  systems  were  not  studied  carefully. 
The  excretory  system  is  made  up  of  coiling  trunks  and  anastomosing 
branches  in  the  head  and  extreme  anterior  neck  region.  In  the  pro- 
glottids  two  pairs  of  main  lateral  excretory  vessels  can  be  made  out. 

The  sexual  organs  (Fig.  102)  in  their  character  and  relations  are 
typical  of  the  snake  Proteocephalids.  The  genital  sinus  is  marginal, 
irregularly  alternating  and  situated  near  the  middle,  or  it  may  be  as 
far  anterior  as  the  end  of  the  first  %  of  the  proglottid.  It  is  marked 
by  a  slight  elevation  which  becomes  more  pronounced  in  the  more  elon- 
gated proglottids.  The  vagina  and  cirrus  open  into  the  common  shallow 
genital  sinus.  The  vagina  lies  either  anterior  or  posterior  to  the  cirrus- 
pouch  but  more  frequently  anterior.  Rarely  the  vagina  is  dorsal  to  the 
cirrus-sheath. 

The  testes  are  very  numerous,  150-215.  They  are  arranged  in  two 
fields,  leaving  a  free  median  zone.  In  elongated  mature  and  ripe  pro- 
glottids the  testes  are  oval  or  even  polygonal.  They  measure  as  much 
as  0.053  by  0.106  mm.  It  is  to  be  noted  that  the  testes  tend  to  occur  in 
groups  of  2,  3  or  4.  In  these  cases  the  limiting  membranes  of  the  testes 
touch  each  other.  The  vasa  efferentia  may  be  clearly  seen  in  some  toto 
preparations.  They  much  resemble  the  vasa  efferentia  in  other  species. 
The  vas  deferens  in  young  proglottids  is  nearly  straight.  It  arises  in 
the  middle  of  the  proglottid  and  extends  to  the  cirrus-pouch  which  it 
enters.  As  the  proglottid  becomes  older  the  vas  deferens  becomes  longer 
and  more  and  more  coiled  until  in  mature  and  ripe  proglottids  it  forms 
a  heavy  mass  of  colls  lying  between  the  cirrus-pouch  and  the  middle  of 
the  segment.  When  filled  with  deeply  stained  spermatozoa  this  mass 
becomes  very  prominent.  Upon  entering  the  cirrus-pouch  the  vas 
deferens  becomes  the  ductus  ejaculatorius,  which  is  coiled  several  times 
before  passing  over  into  the  heavier-walled  cirrus.  The  cirrus  is  some- 
what swollen  but  is  much  smaller  than  in  Crepidohothrium  gerrardii, 
0.  calmettei,  or  0.  grandis.  When  protruded  it  is  slender  and  of  even 
diameter  from  tip  to  base.  Its  length  protruded  is  about  0.200  mm. 
This  measurement  is  taken    from  a  specimen    from    Oklahoma.     The 


207]  PROTEOCEPHALIDAE—LA  RUE  207 

cirrus-pouch  is  more  slender  than  in  several  other  species  of  Ophiotae- 
nia.  In  mature  and  ripe  proglottids  it  measures  from  0.255-0,320  mm. 
long  by  0.08-0.09  mm.  broad.  In  maturing  proglottids  it  measures 
about  0.25  mm.  long  and  is  considerably  narrower  than  in  ripe  ones. 
Its  length  goes  into  the  proglottid  breadth  3-4  times.  In  mature  and 
ripe  proglottids  the  vagina  very  near  its  opening  to  the  exterior  has  a 
wide  but  short  inflated  region.  A  sphincter  vaginae  has  been  found. 
In  this  region  the  vagina  is  richly  set  with  gland  cells.  In  its  course 
to  the  interovarial  space  the  vagina  bends  somewhat  anteriad  and  then 
inward  and  posteriad,  crossing  the  numerous  coils  of  the  vas  deferens 
but  not  crossing  the  cirrus-pouch.  Just  before  the  ovary  is  reached  the 
vagina  is  somewhat  twisted  or  even  thrown  into  small  coils.  A  small 
receptaculum  seminis  can  be  seen  in  some  preparations.  The  ovarian 
lobes  are  long  flattened  bodies  of  irregular  outline.  They  are  not  solid 
but  are  made  up  of  short  heavy  anastomosing  tubes  of  irregular  shape. 
Just  back  of  the  mid-piece  of  the  ovary  the  muscular  oocapt  can  be 
noted  even  in  toto  mounts.  The  coils  and  connections  of  the  oviduct, 
vitelline  duct,  uterine  passage  and  lower  vagina  cannot  be  made  out 
distinctly.  An  ootype  and  shell-gland  have  been  seen  in  some  prepara- 
tions. 

The  vitellaria  are  loosely  follicular  and  lateral.  The  paired  vitel- 
line ducts  cross  the  ovarian  lobes.  The  uterus  in  maturing  proglottids 
is  a  median  tube  from  which  as  the  proglottid  becomes  older  lateral 
pouches  arise  by  the  method  that  has  been  described  for  0.  filaroides 
by  La  Rue  (1909).  There  are  20-30  large  pouches  on  either  side  and 
a  number  of  smaller  ones.  No  uterine  pores  have  been  seen.  In  one 
toto  preparation  the  uterine  passage  may  be  traced  to  a  point  0.29  mm. 
anterior  to  the  mid-piece  of  the  ovary  where  it  discharges  into  the 
uterus. 

The  eggs  are  furnished  with  three  membranes,  an  outer  one  which 
is  smooth,  thin  and  hyaline,  a  middle  membrane  thick  and  granular, 
and  an  inner  thin  membrane  surrounding  the  embryo.  The  outer 
spheroidal  membrane  measures  0.045-0.06-0,100  mm.  and  the  embryos 
are  0.018-0.021  mm.  in  diameter.  Plerocercoids  were  found  encysted  in 
the  intestine  and  liver  of  the  host  caught  at  Havana,  Illinois. 

This  species  is  much  smaller  than  Crepidohothrium  gerrardii 
(Baird),  0.  marenzelleri  (Barrois),  0.  calmettei  (Barrois)  and  0.  tri- 
meresuri  (Parona)  in  most  respects  tho  it  may  exceed  some  of  these  in 
length.  It  most  closely  resembles  0.  nattereri  (Parona),  likewise  from 
one  of  the  Colubrinae.  It  has  a  larger  head  and  larger  proglottids  and 
a  greater  number  of  testes  than  0,  nattereri.  0.  perspicua  is  much 
larger  than  0.  filaroides  La  Rue  from  Amhlystoma  tigrinum  Green,    Its 


208  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [208 

head  is  smaller,  its  testes  are  more  numerous  and  the  position  of  its 
genital  pore  is  very  different  from  that  of  0.  filaroides.  0.  perspicua 
differs  from  0.  lonnhergii  (Fuhrmann)  from  Necturus  maculosus  Raf. 
in  having  a  smaller  head  and  smaller  suckers  and  more  numerous  testes. 


OPHIOTAENIA  LACTEA  (Leidy)  Sp.  Inq. 

1855:     Taenia  lactea  Leidy  1855:443 

1911 :     Ophiotaenia  lactea  La  Rue  1911 :481 

On  account  of  the  inaccessibility  of  Leidy 's  description,  it  is  here 
quoted  in  full: 

"Taenia  lactea  Leidy. — Head  small,  continuous  with  the  neck,  without  ros- 
tellum ;  acetabula  anterior,  hemispherical,  situated  at  the  four  angles.  Neck  mod- 
erately long.  Segments  anteriorly  transversely  oblong,  posteriorly  longer  than  the 
breadth,  square  with  rounded  angles.  Generative  apertures  marginal  (indistinct 
in  the  specimen.) 

"HAB. — One  specimen  i6  inches  long  and  5^  of  a  line  wide  was  found  in 
the  intestine  of  Tropidonotus  sipedon.  In  alcohol  the  specimen  contracted  one- 
half  the  original  length  and  widened  to  i  line." 

La  Rue  (1911:481)  made  this  a  species  inquirenda  in  his  newly 
established  genus  Ophiotaenia.  This  is  probably  a  species  of  Ophiotae- 
nia and  may  be  the  same  as  the  species  reported  by  the  writer  from 
Natrix  rhomhifer.  Unfortunately  the  writer  has  had  as  yet  no  oppor- 
tunity to  make  a  study  of  any  Ophiotaenia  from  Natrix  sipedon  and 
hence  is  unable  to  form  an  opinion  of  the  likelihood  that  the  two  species 
of  host  harbor  the  same  parasites.  Leidy 's  description  is  too  inadequate 
to  permit  a  careful  determination  of  the  position  of  his  form  and  since 
his  specimens  could  not  be  had  by  the  writer  for  study  his  species  is 
considered  to  be  a  species  inquirenda.  Further  research  may  indicate 
its  proper  position. 


209]  PROTEOCEPHAUDAE—LA  RUE  209 

OPHIOTAENIA  FILAROIDES  La  Rue 

[Figs.  26-28,  43-46,  103-105] 

1909:    Proteocephalus  filaroides  La  Eue  1909:17-49 

1911 :     Ophiotaenia  filaroides  La  Rue  1911 :481 

Specific  Diagnosis:  Characters  of  genus.  Worms  attenuate,  small, 
thin,  flat.  Length  80-110  mm.,  maximum  breadth  about  0.80-0.90  mm. 
Color  white,  in  life  somewhat  translucent.  Strobilation  not  evident. 
Intersegmental  furrows  shallow.  Surface  smooth.  Scolex  globose, 
flattened  dorsoventrally,  with  conical  apex,  without  apical  depression 
or  fifth  sucker.  No  rostellum.  No  spines.  Head  not  marked  by  fur- 
rows. Breadth  of  head  0.366-0.46  mm.  Suckers  deep,  muscular,  oval  in 
outline,  maximum  dimension  0.165-0.184  mm.  Neck  narrow,  3-4  mm. 
long.  First  proglottids  0.30-0.36  mm.  broad  by  0.10-0.17  mm.  long. 
Mature  proglottids  quadrate  or  longer  than  broad.  Ripe  proglottids 
from  1.6  mm.  long  by  0.8  mm.  broad  to  4.0  mm.  long  by  0.75  mm.  broad. 
End-proglottid  present.  Musculature  weak.  Parenchyma  coarse,  filled 
with  large  fat  globules. 

Genital  organs  typical  of  genus.  No  genital  papilla.  Genital  pore 
marginal,  irregularly  alternating,  situated  at  end  of  first  fifth  of  pro- 
glottid. Testes  0.05-0.06  mm.  in  diameter,  70-114  in  number,  arranged 
in  two  lateral  fields.  Ductus  ejaculatorius  with  a  few  coils  in  cirrus- 
pouch.  Cirrus  weakly  muscled,  slender,  cylindrical,  0.2-0.3  mm.  long 
when  protruded.  Cirrus-pouch  about  0.22  mm.  long  by  0.11  mm.  broad. 
Vagina  always  anterior  to  cirrus-pouch,  not  crossing  latter.  Weak 
sphincter  vaginae  and  small  receptaculum  seminis  present.  Ovarian 
lobes  thin,  alate,  composed  of  anastomosing  tubules.  Organs  of  inter- 
ovarial  space  typical  of  genus.  Vitellaria  with  large  follicles.  Uterus, 
when  developed,  with  25-35  lateral  pouches  on  either  side.  Uterine 
pores  8-12.  Eggs  with  three  membranes.  Outer  one  0,035-0,100  mm,, 
second,  0.030  mm.,  embryo  0.021  mm.  in  diameter. 

Habitat:  Intestine  and  rectum  of  Amhly stoma  tigrinum  (Green) 
[type  host],  Nebraska  (type  locality)  and  Kansas. 

Type:  Unnumbered  alcoholics  in  collection  of  La  Rue  and  slides 
of  same.    Autotype  in  collection  of  Dr.  H.  B,  Ward. 

This  species  was  first  described  by  La  Rue  (1909:17-49)  as  Proteo- 
cephalus filaroides.  So  far  as  the  writer  has  been  able  to  find  out  no 
one  else  has  worked  on  the  species.  La  Rue  (1911:481)  included  this 
species  in  a  list  of  species  of  Ophiotaenia. 


210  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [210 

This  material  was  obtained  from  Amblystoma  tigrinum  (Green) 
caught  in  ponds  in  Nebraska,  both  at  Crete  and  in  Cherry  County,  also 
in  a  pond  at  Belleville,  Kansas.  The  specimens  were  taken  during  the 
summers  of  1907  and  1908,  The  data  presented  here  are  for  the  most 
part  extracted  from  the  former  article  (La  Eue,  1909)  but  the  data 
regarding  the  character  of  the  fifth  sucker  are  new. 

These  cestodes  are  attenuate,  thin  and  flat.  In  life  they  are  white, 
at  times  somewhat  translucent.  The  chain  presents  no  evident  strobi- 
lation,  for  the  proglottids  are  closely  joined  by  their  full  breadth  and 
the  intersegmental  furrows  are  shallow.  There  are  no  longitudinal  or 
transverse  furrows  in  preserved  specimens.  The  strobila  measures  80- 
110  mm.  in  length  by  a  maximum  breadth  of  0.80-0.90  mm.  The  scolex 
(Fig.  26)  is  globose,  flattened  somewhat  dorsoventrally,  and  possessing 
a  smooth  conical  apex  in  which  there  is  no  depression  nor  fifth  sucker. 
There  is  no  rostellum  and  no  spines.  The  surface  of  the  head  is  usually 
smooth  and  but  rarely  marked  by  furrows.  The  head  measures  0.366- 
0.40  mm.  broad.  Four  oval  suckers  are  borne  at  the  broadest  part  of 
the  head.  They  measure  0.165-0.184  mm.  in  the  maximum  dimension. 
The  suckers  have  deep  cavities  and  their  musculature  is  well  developed. 

As  in  many  other  species  of  this  group  which  possess  no  functional 
fifth  sucker  there  is  in  the  tissue  below  the  surface  of  the  apex  of  the 
head  a  structure  which  the  writer  formerly  called  an  endorgan.  This 
structure  appears  as  a  small  mass  of  tissue  surrounded  by  a  definite 
membrane.  The  mass  of  tissue  contains  a  few  nuclei  and  a  few  scat- 
tered fibers  which  may  be  muscle  fibers.  In  the  adult  this  structure  is 
about  0.063  mm.  long  by  0.034  mm.  broad.  It  has  no  opening  to  the 
exterior.  Two  histological  drawings  of  the  adult  organ  are  to  be  found 
in  a  previous  paper  (La  Rue  1909,  Figs.  13  and  17).  In  the  plerocercus 
this  structure  is  much  larger  than  in  the  adult.  This  fact  was  pointed 
out  in  the  author's  paper  of  1909  and  figures  were  given  to  show  the 
difference  in  size.  These  figures  have  been  reproduced  (Figs.  27,  28). 
If  sections  through  the  head  of  plerocereoids  of  this  species  be  examined 
one  notes  that  the  endorgan  has  many  points  in  common  with  the  func- 
tional fifth  sucker.  There  is  a  sucker  cavity  communicating  with  the 
exterior,  a  basement  membrane,  muscle  fibers  (sub-cuticular  and  radial). 
The  musculature  about  the  organ  is  also  like  that  about  other  suckers. 
These  histological  features  are  shown  in  drawings  which  are  reproduced 
(Figs.  43-46).  This  structure  is,  however,  undergoing  a  marked  modi- 
fication. The  sucker  cavity  is  completely  or  partially  filled  with  a  gran- 
ular mass  of  apparently  the  same  texture  as  that  which  makes  up  the 
greater  bulk  of  the  sucker  itself.  The  radial  muscles  are  no  longer 
arranged  in  such  beautiful  order  as  in  other  suckers  but  they  seem  to 


211]  PROTEOCEPHALIDAE—LA  RUE  211 

be  twisted  and  pressed  out  of  the  regular  position  and  spread  far  apart 
by  the  granular  mass.  Radial  muscle  fibers  showing  the  most  typical 
arrangement  are  to  be  seen  in  figure  45.  Nuclei  are  plainly  seen  and 
these  are  figured  as  the  larger  oval  black  spots.  The  drawing  of  the 
endorgan  (Fig.  43)  is  made  from  a  head  cut  in  a  frontal  or  sagittal 
plane.  It  shows  the  opening  to  the  exterior,  the  basement  membrane 
and  the  cuticular  lining  of  a  part  of  this  opening.  It  also  shows  the 
cut  ends  of  the  muscle  fibers  next  to  the  basement  membrane.  Some 
of  these  same  features  may  be  well  seen  in  other  drawings  (Figs.  45 
and  46)  which  are  drawn  from  transverse  sections  of  other  heads. 

It  is  evident  from  the  structures  here  described  and  the  drawings 
which  illustrate  them  that  this  organ  is  in  reality  a  sucker.  It  is  a 
sucker  which  evidently  comes  to  a  certain  state  of  development  in  which 
as  pointed  out  by  the  writer  in  his  former  paper  on  this  species  (La 
Rue  1909:25),  it  is  larger  than  the  other  suckers.  The  relative  sizes 
of  this  organ  are  shown  in  drawings  reproduced  (Figs.  27,  28)  from 
the  former  paper.  This  enlargement  is  due  to  a  hypertrophy  charac- 
terized by  the  presence  of  granules.  Altho  the  stages  succeeding  this 
hypertrophy  have  not  been  followed  out  it  is  plain  that  the  hypertrophy 
is  succeeded  by  an  atrophy  of  the  tissues.  Since  the  granules  are  not 
present  in  the  adult  organ  they  must  disappear  either  as  a  result  of 
streaming  out  of  the  sucker  opening  before  that  is  closed  or  they  may 
be  absorbed  by  the  organism.  The  sucker  opening  and  the  sucker  cavity 
and  all  traces  of  the  cutieula,  outer  basement  membrane,  and  muscles 
about  the  sucker  cavity  are  obliterated  by  the  time  this  organ  is  found 
in  the  adult  cestode  head.  Thus  the  sucker  loses  all  connection  with 
the  exterior.  It  retains  its  limiting  basement  membrane,  some  of  its 
nuclei  and  perhaps  a  few  scattered  muscle  fibers.  The  conclusion  is 
then  that  this  endorgan,  or  muscle-plug  as  Johnston  (1909  et  seq.)  has 
called  it,  is  a  vestigial  fifth  sucker.  That  the  writer  was  dealing  with 
the  plerocercoids  of  this  species  is  attested  by  the  facts  brought  out  by 
his  feeding  experiments  (La  Rue  1909)  in  which  he  fed  plerocercoids 
taken  from  the  flesh  of  Amblystoma  to  other  uninfected  Amblystoma 
and  the  latter  became  heavily  infected  with  the  cestode  Ophiotaenia 
filaroides. 

The  narrow  neck  is  3-4  mm.  long.  This  is  followed  by  a  region  of 
short  proglottids  which  gradually  become  longer  and  broader.  The 
increase  in  length  is  more  rapid  than  the  increase  in  breadth  hence  the 
proglottids  change  progressively  from  broader  than  long  to  quadrate 
and  then  to  longer  than  broad.  The  youngest  proglottids  measure  0.30- 
0.36  mm.  broad  by  0.10-0.17  mm.  long.  Ripe  proglottids  measure  1.6 
mm.  long  by  0.8  mm.  broad  and  in  some  cases  as  much  as  4.0  mm.  long 


212  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [212 

by  0.75  mm.  broad.  An  end-proglottid  with  a  rounded  posterior  end 
may  be  present,  but  it  is  lost  with  the  first  ripe  proglottids. 

The  musculature  of  the  body  is  weakly  developed.  This  is  particu- 
larly true  of  the  longitudinal  muscles  which  lie  between  the  cortical  and 
medullary  layers  of  the  body.  In  the  head  the  excretory  system  is 
made  up  of  coils  of  small  anastomosing  vessels  which  are  connected 
with  the  two  pairs  of  main  lateral  vessels  extending  through  the  length 
of  the  strobila.  In  the  strobila  both  pairs  of  lateral  vessels  lie  within 
the  dermo-muscular  sheath.  The  genital  canals,  cirrus-pouch  and 
vagina,  always  pass  between  the  dorsal  and  ventral  excretory  vessels. 
The  latter  are  connected  with  the  exterior  by  numerous  branches  which 
discharge  on  the  dorsal  or  ventral  surface  more  frequently  than  on  the 
margin  of  the  segment.  Regularly  situated  foramina  secundaria  do  not 
exist.  A  transverse  excretory  commissure  uniting  the  ventral  vessels  in 
the  posterior  part  of  the  proglottid  has  been  observed.  The  meshes  of 
the  parenchyma  are  coarse,  forming  numerous  large  spaces.  In  mate- 
rial prepared  by  ordinary  methods  this  tissue  appears  to  be  nothing 
more  than  abnormally  loose  parenchyma.  When  fresh  material  is 
stained  with  ''Sudan  III"  or  is  treated  with  osmic  acid  these  spaces 
are  found  to  be  filled  with  large  fat  globules.  These  tests  have  likewise 
been  applied  by  the  writer  to  pieces  of  0.  lonnhergii  with  similar  results. 
Fat  globules  have  been  found  in  the  tissues  of  species  of  Proteocephalus 
but  in  this  case  the  fat  globules  are  small.  Tests  have  also  been  made 
on  Taenia  saginata  (?),  Taenia  serrata,  and  Dipylidium  caninum  and 
in  each  case  fats  have  been  found.  Tests  for  fat  have  not  been  made 
on  the  cestodes  of  snakes  but  all  the  species  examined  show  the  loose 
parenchyma  with  large  spaces,  which  the  fat  globules  once  occupied. 

The  genital  organs  (Figs.  103,  104,  105)  are  typical  of  the  genus. 
A  genital  papilla  is  not  present.  The  genital  pore  is  marginal,  irregu- 
larly alternating,  and  situated  near  the  end  of  the  anterior  fifth  of  the 
proglottid.  Through  the  genital  pore  the  small  genital  atrium  is  con- 
nected with  the  exterior.  Into  the  genital  atrium  both  cirrus  and  vagina 
open.  The  vagina  always  lies  anterior  to  the  cirrus-pouch.  The  testes 
(Fig.  105)  measure  about  0.05-0.06  mm.  in  diameter.  They  number 
about  70-114  and  they  are  situated  in  two  broad  fields  on  either  side  of 
a  free  median  zone.  They  occupy  a  position  dorsal  to  the  uterus.  A 
much  coiled  vas  deferens  forms  a  mass  which  extends  from  the  cirrus- 
pouch  to  the  midfield  of  the  proglottid.  This  mass  of  coils  functions  as 
a  vesicula  seminalis.  Upon  entering  the  cirrus-pouch  the  vas  deferens 
becomes  the  ductus  ejaculatorius  which  makes  1-3  coils  before  passing 
over  into  the  straight  and  weakly  muscled  cirrus.  When  protruded 
the  cirrus  measures  0.2-0.3  mm.  in  length.    It  is  slender  and  of  almost 


213]  PROTEOCEPHALIDAE—LA  RUE  213 

uniform  diameter  from  base  to  tip.  The  cirrus-pouch  is  weakly  muscled. 
Its  length  is  about  0.22  mm.  and  its  breadth  about  0.11  mm.,  being 
broadest  at  its  inner  end. 

The  vagina  (Fig.  105)  always  lies  anterior  to  the  cirrus-pouch  and 
does  not  cross  the  latter.  There  are  no  coils  of  vagina  anterior  to  the 
interovarial  space.  A  weak  sphincter  vaginae  and  a  small  receptacu- 
lum  seminis  are  present.  The  lobes  of  the  ovary  are  thin  and  somewhat 
alate  in  shape.  They  are  made  up  of  anastomosing  tubules.  The  organs 
(Fig.  104)  of  the  interovarial  space  are  typical  of  the  genus.  Vitelline 
follicles  are  large  and  are  arranged  in  the  lateral  fields  as  in  the  other 
members  of  the  group.  The  uterus  in  mature  proglottids  is  a  median 
tube.  From  this  tube  there  arise  both  lateral  and  ventral  diverticula 
after  the  manner  described  by  La  Rue  (1909:33-37).  The  lateral 
diverticula  in  fully  ripe  proglottids  extend  to  the  vitellaria  which  by 
this  time  have  degenerated  to  a  large  extent.  They  number  from  25  to  35 
on  either  side.  The  ventral  diverticula  number  about  8-12.  These  are 
short  and  usually  pointed.  In  time  they  pierce  the  ventral  body  wall 
forming  the  uterine  pores.  The  ventral  body  wall  now  gives  way  along 
this  line  of  perforations  thus  causing  a  rift  which  extends  from  one 
end  of  the  proglottid  to  the  other.  Through  this  rift  the  eggs  are  dis- 
charged. The  eggs  are  covered  with  three  membranes  of  the  character 
usual  to  the  group.  The  outer  thin  and  hyaline  membrane  varies  in 
diameter  from  0.035  to  0.10  mm.  This  variation  is  due  in  part  to  the  fact 
that  when  it  comes  in  contact  with  the  water  the  outer  membrane 
swells  up  greatly.  The  second  membrane  is  thick  and  granular,  about 
0.030  mm.  in  diameter.  The  inner  membrane  which  closely  invests  the 
embryo  is  thin.  The  six-hooked  embryo  measures  about  0.021  mm.  in 
diameter.  The  larval  form  of  this  cestode  is  frequently  found  encysted 
in  the  abdominal  viscera  and  body  muscles  of  the  host,  Amhlystoma 
tigrinum.  They  have  also  been  found  free  in  the  body  cavity  of  the 
same  host.    As  yet  they  have  not  been  found  in  an  invertebrate  host. 

Altho  this  species  shows  marked  similarities  with  some  of  the  spe- 
cies oF  cestodes  infesting  snakes  it  most  nearly  resembles  Ophiotaenia 
lonnhergii  (Fuhrmann)  which  infests  the  amphibian,  Necturus  maculo- 
sus  Raf.  It  differs  from  this  species  in  size  relations,  in  the  character 
of  the  excretory  vessels,  and  in  the  relations  of  the  cirrus-pouch  to  the 
vagina.  There  are  some  minor  points  in  which  the  two  species  from 
the  amphibia  differ  from  the  species  infesting  snakes  but  as  yet  charac- 
ters have  not  been  found  by  which  they  can  be  separated  from  the 
genus  Ophiotaenia.  These  two  species  certainly  do  not  belong  with  the 
genus  Proteocephalus. 


214  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [214 

OPHIOTAENIA  LONNBERGII    (Fuhrraann) 

[Figs.  119,  189] 

1895:    Ichthyotaenia  lonnhergii      Fuhrmann  1895:218-226 

1909:    Proteocephalus  lonnhergii     La  Rue  1909:43 

1911 :     Ophiotaenia  lonnhergii  La  Rue  1911 :481 

Specific  Diagnosis:  Characters  of  the  genus.  Length  as  much  as 
17-19  cm.  Breadth  as  much  as  1.35  mm.  Strobilation  usually  indis- 
tinct. Intersegmental  furrows  shallow.  Proglottids  attached  by  full 
width.  Scolex  globose,  flattened  dorsoventrally,  0.50-0.60  mm.  in 
breadth.  No  rostellum,  no  spines,  no  functional  fifth  sucker.  Suckers 
prominent,  muscular,  oval  or  round,  deep,  measuring  0.24-0.26  mm. 
long  by  0.14-0.22  mm.  broad.  Opening  in  oval  suckers  0,20  by  0.10 
mm.  Neck  0.375-0.54  mm.  in  minimum  breadth,  length  about  2.0  mm. 
First  proglottids  about  0.5  mm.  long  by  0.50  mm.  broad.  Mature  pro- 
glottids quadrate,  0.85-1.0  mm.  square  or  longer  than  broad,  measuring 
as  much  as  2.5  mm.  long  by  0.45-0.5  mm.  broad.  Ripe  proglottids  not 
observed.  Two  pairs  of  main  lateral  excretory  vessels  situated  at  some 
distance  from  lateral  margins  of  strobila.  Vessels  spiral  in  form. 
Transverse  commissure  present  in  posterior  end  of  each  proglottid. 
Many  branches  extending  from  dorsal  and  ventral  vessels  to  exterior. 
In  head  many  branches  of  anastomosing  vessels. 

Genital  pore  marginal,  irregularly  alternating,  situated  at  end  of 
first  one-third  or  two-fifths  of  proglottid.  Testes  90-160  in  number, 
round  or  oval,  0.05-0.08-0.12  mm.  in  maximum  dimension,  situated  in 
two  lateral  fields  extending  laterad  of  excretory  vessels.  Midzone  of 
proglottid  not  entirely  free  of  testes.  Vas  deferens  forming  a  large 
mass  of  coils  posterior  to  cirrus-pouch.  Cirrus-pouch  0.185-0.280  mm. 
long  by  0.05-0.085-0.10  mm.  broad.  Ductus  ejaculatorius  in  several 
coils,  frequently  forming  a  small  vesicula.  Cirrus  not  muscular,  when 
protruded  cylindrical  and  0.13-0.15  mm.  long  by  0.045-0.05  mm.  thick. 
Vagina  anterior  or  posterior  to  cirrus-pouch.  Vaginal  opening  some- 
times dorsal  to  latter,  but  vagina  never  crossing  cirrus-pouch.  A  large 
dilatation  in  vagina  of  some  proglottids.  Sphincter  vaginae  and  recep- 
taculum  seminis  present.  Ovary  thin,  flat,  lobes  wedge-shaped  or  alate. 
Organs  of  interovarial  space  typical  of  genus.  Vitellaria  sparse.  Uterus 
in  past  mature  proglottids  possessing  25-40  lateral  pouches.  Eggs  im- 
mature in  specimens  observed.  Yolk-mass  0.010-0.012  mm.  in  diameter. 
Egg  membranes  not  seen.     Embryos  not  yet  developed. 

Habitat:  In  intestine  of  Necturus  maculosus  Rar.  Fuhrmann 
does  not  state  the  locality  of  the  host  from  which  his  type  specimens 


215]  PROTEOCEPHALIDAE—LA  RUE  215 

came.  The  writer  has  frequently  found  the  species  in  Necturus  from 
Ohio  and  Indiana  which  had  been  brought  to  the  laboratory  for  dissec- 
tion, Fuhrmann's  specimen  was  obtained  from  Prof.  F.  Zschokke  who 
received  it  from  Prof.  R.  Burckhardt. 

Type:    Prepared  slides  in  collection  of  Fuhrmann. 

Fuhrmann  (1895:218-226)  described  and  delineated  this  species, 
calling  it  Ichthyotaenia  Idnnhergii.  Without  attempting  a  description 
of  the  species  it  was  referred  to  by  La  Rue  ( 1909 :43 )  as  Proteocephalus 
lonnbergii.  In  a  more  recent  paper  La  Rue  (1911:481)  pointed  out 
that  this  was  to  be  considered  as  a  species  of  a  new  genus,  Ophiotaenia. 

This  study  is  based  on  slides  and  alcoholics  in  the  collection  of 
La  Rue.  The  material  has  been  carefully  compared  with  Fuhrmann's 
preparations  of  the  type  which  Professor  Ward  very  kindly  secured 
for  the  writer's  use.  The  material  was  thus  found  to  be  identical  with 
Fuhrmann's.  Reference  will  be  made  from  time  to  time  to  the  work 
of  Fuhrmann  (1895)  from  which  certain  data  were  secured. 

This  form  is  more  robust  than  0.  filaroides.  It  may  reach  a  length 
of  17.0  cm.  or  more.  Fuhrmann's  specimen,  an  immature  worm,  meas- 
ured 19.0  cm.  The  worm  is  thin  and  flat.  Its  breadth  varies  consider- 
ably. The  maximum  breadth  in  the  writer's  specimens  was  1.275  mm. 
Fuhrmann's  material  had  a  slightly  greater  breadth.  The  strobilation 
is  rarely  distinct.  The  margins  of  the  strobila  are  quite  smooth.  In- 
frequently the  posterior  part  of  the  worm  may  show  a  distinct  strobila- 
tion. The  proglottids  are  attached  by  their  full  width.  Transverse 
folds  are  rare  but  shallow  longitudinal  furrows  are  not  uncommon. 
The  scolex  (Fig.  119)  is  somewhat  globose  and  is  flattened  dorsoven- 
trally.  On  it  are  four  prominent  suckers  situated  on  its  broadest  part. 
In  breadth  the  scolex  varies  from  0.50  to  0.60  mm.  There  is  no  rostel- 
lum,  no  spines  and  no  functional  fifth  sucker.  The  suckers  are  round  or 
oval  in  outline  with  deep  cavities  and  a  strong  musculature.  In  length 
the  suckers  vary  from  0.24  to  0.25  mm.  and  the  breadth  from  0.14  to 
0.22  mm.  The  opening  of  the  more  oval  suckers  measures  about  0.20 
by  0.10  mm.  Immediately  behind  the  head  the  neck  has  a  breadth  of 
0.375-0.54  mm.  The  first  traces  of  segmentation  occur  about  2  mm. 
posterior  to  the  head. 

Young  proglottids,  according  to  Fuhrmann,  measure  0.27  mm,  in 
length.  The  first  discernible  proglottids  in  the  writer's  material  are 
about  0.050  mm.  long  by  0.50  mm.  broad.  The  length  and  breadth  of 
the  proglottids  increase  rapidly  for  a  distance  then  the  length  increases 
while  the  breadth  decreases.  In  this  material  the  maximum  breadth  of 
1.275  mm.  was  reached  long  before  the  proglottids  were  mature.    Pro- 


216  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [216 

glottids  in  this  region  are  1,275  mm.  broad  by  0.30-0.40  mm.  long. 
Mature  proglottids  are  about  0.85-1.0  mm.  square,  or  they  may  be  longer 
than  broad.  Some  of  the  elongated  proglottids  measure  2.5  mm.  long 
by  0.45-0.5  mm.  broad.  In  Fuhrmann's  specimens  the  last  proglottids 
were  0.7  mm.  long  by  1.35  mm.  broad.  Neither  Fuhrmann  nor  the 
writer  has  seen  the  ripe  proglottids.  All  the  hosts  examined  by  the 
latter  had  been  in  captivity  for  some  time  and  were  examined  in  the 
spring  months.  It  may  be  that  the  parasites  attain  the  sexually  ripe 
condition  later  in  the  season. 

The  excretory  system  (Fig.  189)  is  made  up  of  two  pairs  of  main 
lateral  vessels,  dorsal  and  ventral.  These  are  situated  much  farther 
mesad  than  is  usually  the  case  with  this  group.  They  take  a  spiral 
path  through  the  tissues.  From  them  at  frequent  intervals  smaU 
branches  pass  to  the  dorsal  and  ventral  surfaces.  A  transverse  excre- 
tory commissure  is  present  in  the  posterior  part  of  each  segment.  In 
the  head  the  main  excretory  vessels  are  connected  by  means  of  many 
anastomosing  branches.  The  parenchyma  resembles  that  of  0.  fla- 
roides.  Chalkbodies  of  large  size  are  abundant,  especially  in  the  head 
and  neck.    The  musculature  is  weakly  developed. 

The  genital  pore  alternates  irregularly,  is  marginal,  and  is  situated 
at  the  end  of  the  first  third  or  two-fifths  of  the  proglottid.  Cirrus- 
pouch  and  vagina  pass  to  the  exterior  between  the  ventral  and  the 
dorsal  excretory  vessels.  The  male  organs  mature  first.  Testes  number 
from  90  to  160,  the  number  varying  greatly  in  the  proglottids  of  the 
same  strobila.  Elongated  testes  measure  0.05-0.08  and  even  as  much  as 
0.120  mm.  in  maximum  dimension.  The  testes  (Fig.  189)  lie  in  two 
broad  fields  which  extend  laterad  and  mesad  of  the  excretory  vessels. 
In  many  proglottids  an  occasional  testis  lies  in  the  midfield.  The  testes 
are  dorsal  to  the  uterus.  In  mature  proglottids  the  vas  deferens  forms 
a  large  mass  of  coils  which  function  as  a  vesicula  seminalis.  This  mass 
of  coils  is  usually  situated  posterior  to  the  cirrus-pouch.  It  lies  almost 
in  the  mid-field  of  the  segment.  The  cirrus-pouch  is  short  and  broad. 
Its  length  varies  from  0.185  to  0.290  mm.  and  its  breadth  from  0.050  in 
the  case  of  the  long  slender  pouches  to  0.085-0.100  mm.  in  the  shorter 
and  thicker  pouches.  The  cirrus-sheath  is  broadest  at  the  inner  end. 
The  ductus  ejaculatorius  forms  several  coils  and  then  it  passes  over  into 
the  slender  cirrus.  In  some  proglottids  the  ductus  is  dilated  to  form  a 
small  vesicle.  The  cirrus  is  not  muscular.  When  protruded  it  is  cylin- 
drical and  about  0.13-0.15  mm.  long  by  0.045-0.050  mm.  thick.  There 
are  no  coils  of  ductus  ejaculatorius  in  the  base  of  the  protruded  cirrus. 

The  vagina  lies  anterior  or  posterior  to  the  cirrus-pouch  but  its 
opening  is  usually  dorsal  to  the  latter.    When  the  vagina  occupies  the 


217]  PROTEOCEPHALIDAE—LA  RUE  217 

position  anterior  to  the  cirrus-pouch  it  never  crosses  the  latter  but 
passes  beyond  it  and  then  dips  down  below  the  coils  of  the  vas  deferens. 
In  many  mature  proglottids  the  initial  part  of  the  vagina  is  greatly 
dilated  and  this  dilatation  extends  for  some  distance  down  the  length 
of  the  vagina.  There  are  no  coils  of  the  vagina  anterior  to  the  ovary 
but  it  may  lie  in  sinuous  curves.  A  weak  sphincter  vaginae  close  to  the 
external  opening  and  a  small  receptaculum  seminis  near  the  mid-piece 
of  the  ovary  are  present.  The  ovary  varies  in  shape  with  the  state  of 
contraction  of  the  proglottid.  It  is  always  thin  and  flat  and  is  made  up 
of  anastomosing  tubules  as  in  0.  filaroides.  In  very  broad  but  short 
proglottids  the  lobes  of  the  ovary  are  long  slender  wedge-shaped  struc- 
tures the  apices  of  which  lie  in  the  mid-field  of  the  proglottid.  In 
attenuated  proglottids  the  lobes  are  much  shortened  and  broader  than 
in  long  proglottids,  in  which  case  the  ovary  may  be  distinctly  alate  in 
form.  The  organs  of  the  interovarial  space  are  arranged  about  as 
delineated  for  0.  filaroides  (Fig.  104).  The  vitellaria  are  lateral,  com- 
posed of  medium-sized  sparse  follicles.  Since  no  ripe  proglottids  have 
been  seen  a  well  developed  uterus  cannot  be  described.  In  one  proglot- 
tid in  which  a  few  eggs  had  been  passed  into  the  uterus  there  were 
from  25  to  40  lateral  pouches  on  either  side.  No  uterine  pores  have 
been  observed.  No  eggs  with  developed  embryos  could  be  found.  A  few 
eggs  which  could  be  but  poorly  seen  within  the  uterus  of  a  toto  prepara- 
tion showed  yolk-masses  measuring  0.010-0.012  mm.  in  diameter.  The 
egg  membranes  could  not  be  measured.    Fuhrmann  saw  no  eggs. 

This  species  tho  most  closely  related  to  0.  filariodes  can  be  readily 
distinguished  from  the  latter  by  its  larger  strobila,  larger  head,  larger 
suckers  and  larger  testes.  The  excretory  vessels  are  spiral  structures 
in  0.  lonnbergii  but  straight  in  0.  filaroides.  The  ovarian  lobes  are 
also  different  in  size  and  shape.  0.  lonnbergii  differs  from  the  Ophio- 
taenia  parasitic  in  snakes  by  the  position  of  its  genital  pore  and  by 
the  less  muscular  character  of  its  cirrus.  Its  ovary  is  also  of  a  different 
shape.  The  position  of  the  excretory  ducts  is  much  farther  mesad  in 
0.  lonnbergii  than  in  any  species  of  Ophiotaenia  at  present  known  from 
snakes. 

While  further  investigation  may  prove  that  the  species  of  Proteo- 
eephalids  infesting  Amphibia  are  generically  different  from  the  species 
infesting  snakes  there  is  at  present  no  justification  in  erecting  a  new 
genus  for  them.  Their  nearest  allies  are  among  the  Ophiotaenia,  hence 
the  writer  places  them  in  that  genus. 


218  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [218 

OPHIOTAENIA  NATTERERI  (Parona) 
[Figs.  139,  194,  195] 


1901 

1908 
1911 


Ichthyotaenia  nattereri        Parona  1901 :4-6 

Ichthyotaenia  nattereri        Schwarz         1908:17-19 
Ophiotaenia  nattereri  La  Rue  1911 :481 


Specific  Diagnosis:  Characters  of  genus.  Length  75-250  mm,, 
maximum  breadth  not  over  1  mm.  Scblex  unarmed,  rounded  but  not 
spheroidal,  not  easily  distinguished  from  the  neck.  Diameter  of  scolex 
0.240-0.250  mm.  Suckers  four  in  number,  circular,  cavity  deep,  margin 
thick,  diameter  of  suckers  0.120-0.150  mm.  No  rostellum  and  no  fifth 
sucker.  Neck  long,  7-8  times  the  length  of  the  head.  First  segments 
broader  than  long,  more  or  less  indistinct.  Mature  proglottids  nearly 
quadrate.  Ripe  proglottids  longer  than  broad,  length  of  same  up  to  2.0 
mm.,  breadth  0.33-0.50  mm.  Last  proglottids  readily  detached.  Seg- 
mentation distinct. 

Genital  aperture  irregularly  alternating,  situated  somewhat  ante- 
rior to  middle  of  proglottid  margin.  Testes  80-100,  0.042  mm.  in  diame- 
ter, situated  in  two  lateral  fields.  Vas  deferens  a  thick  mass  of  coils 
extending  even  past  the  middle  of  the  proglottid.  Ductus  ejaculatorius 
much  coiled.  Protruded  cirrus  slender,  heavy  at  base,  about  0.20  mm. 
long.  Cirrus-pouch  about  0.265-0.280  mm.  long,  extending  from  Vs-'/t 
across  proglottid  breadth.  Vagina  anterior  or  posterior  to  cirrus-pouch. 
Sphincter  vaginae  present.  Uterus,  when  fully  developed,  possessing 
15-20  lateral  pouches  on  either  side.  Eggs  provided  with  three  mem- 
branes. Outer  membrane  covered  with  fine  booklets.  Eggs  0.024  mm. 
in  diameter. 

Habitat:  In  intestine  of  Coluber  sp.  from  Liguria,  a  district  of 
Italy.    The  material  was  collected  by  Parona  in  December,  1897. 

This  species  was  first  described  by  Parona  (1901:4-6)  but  the  de- 
scription was  not  accompanied  by  drawings.  Schwarz  (1908:17-19) 
delineated  and  redescribed  the  species,  using  Parona 's  specimens.  La 
Rue  (1911:481)  included  this  form  in  a  list  of  species  of  Ophiotaenia. 
The  data  upon  which  this  description  is  based  were  secured  from  the 
descriptions  of  Parona  (1901)  and  Schwarz  (1908),  and  from  observa- 
tions made  upon  slides  prepared  from  type  material  which  Professor 
H.  B.  Ward  secured  from  Professor  Parona. 


219]  PROTEOCEPHALIDAE—LA  RUE  219 

This  cestode  is  thin  and  slender.  Its  observed  length  varies  from 
75-250  mm.  and  its  maximum  breadth  is  not  over  1  mm.  The  segmenta- 
tion is  distinct  in  the  posterior  region.  The  single  proglottids  are  thin, 
and  somewhat  translucent.  The  first  proglottids  are  broader  than  long 
and  their  boundaries  between  segments  are  not  distinct.  More  mature 
proglottids  are  nearly  quadrate  while  ripe  proglottids  are  longer  than 
broad.  The  length  of  ripe  proglottids  may  be  as  much  as  2.0  mm.  and 
the  breadth  0.33-0.50  mm.  The  last  proglottids  are  easily  detached 
from  the  chain.  The  head  is  small  and  not  clearly  set  oif  from  the 
neck.  It  is  unarmed,  rounded  somewhat  but  not  spherical ;  its  diameter 
is  0.24-0.25  mm.  It  bears  four  circular  suckers  which  have  a  deep  cavity 
and  a  thick  muscular  wall.  The  diameter  of  the  suckers  varies  from 
0.120  to  0.150  mm.  There  is  no  rostellum  and  no  fifth  sucker.  The  neck 
is  seven  or  eight  times  as  long  as  the  head.  Cuticula,  musculature, 
nervous  system  and  excretory  system  are  very  similar  in  arrangement 
and  character  to  the  same  systems  in  other  species  of  the  genus. 

The  early  developing  sex  organs  may  be  seen  in  quadrate  proglot- 
tids. The  genital  aperture  lies  somewhat  anterior  to  the  middle  of  the 
margin  of  the  segment  and  it  irregularly  alternates  from  right  to  left. 
Schwarz's  figure  of  the  ripe  proglottid  showing  the  main  features  of 
the  reproductive  systems  is  reproduced  (Fig.  195).  The  testes  lie  in 
two  fields  between  the  vitellaria,  thus  leaving  the  median  zone  of  the 
proglottid  free  of  them.  They  number  80-100  and  measure  0.042  mm. 
The  vas  deferens  forms  a  thick  heavy  coil  extending  from  the  cirrus- 
pouch  even  past  the  middle  of  the  proglottid.  Within  the  cirrus-pouch 
there  is  a  very  complicated  mass  of  coils  of  ductus  ejaculatorius.  This 
is  much  more  highly  developed  than  in  any  other  known  species  of 
Ophiotaenia.  The  cirrus  when  protruded  is  heavy  at  the  base,  more 
slender  at  the  tip  and  about  0.200-0.210  mm.  long.  In  preparations 
where  it  was  incompletely  protruded  it  measured  0.108  mm.  long.  The 
cirrus-pouch  is  0.265-0.280  mm.  long.  In  quadrate  and  oblong  proglot- 
tids its  length  goes  into  the  proglottid  breadth  from  3  to  3.5  times. 

The  vagina  opens  into  the  common  genital  pore.  It  may  lie  either 
anterior  or  posterior  to  the  cirrus-pouch.  In  12  out  of  18  proglottids 
examined  by  the  writer  the  vagina  had  a  posterior  position.  At  the 
opening  of  the  vagina  a  sphincter  of  good  size  is  present.  The  course 
of  the  vagina  is  nearly  direct  to  the  middle  of  the  segment,  frequently 
crossing  the  inner  end  of  the  cirrus-pouch  or  even  lying  above  the  cirrus- 
pouch  for  its  full  length.  From  the  middle  of  the  segment  its  course  is 
directly  posteriad  with  sometimes  a  single  loose  coil  just  above  or  ante- 
rior to  the  ovaries.  In  the  interovarial  space  are  the  usual  coils  of  the 
vagina,  oviduct,  uterine  passage,  and  unpaired  vitelline  duct;  here  also 


220  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [220 

are  the  oocapt,  ootype,  and  shell-gland.  The  uterus  appears  in  mature 
proglottids  as  a  median  tube  from  which  later  15-20  lateral  pouches 
arise  on  either  side.  The  ovarian  lobes  are  fairly  dense  compact  masses. 
In  many  proglottids  the  ovary  has  a  more  elongated  and  pointed  form 
than  Schwarz  delineates  it.  The  vitellaria  are  lateral,  follicular  masses, 
reaching  from  the  anterior  end  nearly  to  the  posterior  end  of  the  seg- 
ment. According  to  Schwarz  the  eggs  have  three  membranes.  The 
diameter  of  the  ^gg  is  0.024  mm.  The  outer  membrane  of  ripe  eggs  is 
covered  with  fine  booklets  or  processes  which  terminate  in  small  knobs. 
The  writer  was  unable  to  observe  these  processes  in  his  toto  mounts. 
Schwarz 's  figure  of  the  Qgg  is  reproduced  (Fig.  194). 

This  species  is  easily  differentiated  from  Crepidobothrium  gerrardii, 
0.  marenzelleri,  0.  calmettei,  0.  racemosa,  0.  trimeresuri  and  0.  grandis 
by  its  much  smaller  size,  and  from  0.  perspicua  by  its  slightly  smaller 
head  and  suckers,  by  its  lesser  number  of  smaller  testes,  its  more  volumi- 
nous cirrus-pouch  and  its  lesser  number  of  uterine  pouches.  In  many 
respects  0.  nattereri  and  0.  perspicua  are  quite  similar.  0.  nattereri 
has  a  smaller  head  and  smaller  suckers  than  0.  pigmentata.  Further 
comparisons  with  0.  pigmentata  can  not  be  made  because  of  the  incom- 
pleteness of  the  data  on  the  latter. 


OPHIOTAENIA  RACEMOSA  (Rudolphi; 
[Figs.  140,  191] 

1819:  Taenia  racemosa 

(?)1819:  Taenia  cpluhfi 

1845:  Taenia  racemosa 

1850:  Taenia  racemosa 

1898:  Ichthyotaenia  racemosa 

1898 :  Ichthyotaenia  racemosa 

1908:  Ichthyotaenia  racemosa 

1911 :  Ophiotaenia  racemosa 

Specific  Diagnosis:  Characters  of  genus.  Length  of  strobila  as 
much  as  160  mm.,  breadth  about  1  mm.  Length  of  ripe  proglottids 
about  2  mm.,  breadth  1  mm.  Scolex  easily  distinguishable  from  strobila, 
breadth  about  0.54  mm.  Suckers  four,  nearly  circular,  0.27-0.30  mm. 
in  diameter.  Genital  organs  as  in  genus.  Testes  about  100-120  in 
number,  0.072-0.078  mm.  in  diameter,  located  in  two  broad  fields  which 
tend  to  fuse  at  middle.  Ductus  ejaculatorius  sinuous.  Cirrus-pouch 
long,  reaching  nearly  to  middle  of  proglottid.    Vagina  anterior  or  pos- 


Rudolphi 

1819  :692 

Rudolphi 

1819  :709 

Dujardin 

1845 :610 

Diesing 

1850:511,  in  part 

Barrois 

1898:3 

Liihe 

1898 :652 

Schwarz 

1908 :28-29 

La  Rue 

1911 :481 

221] 


PROTEOCEPHALIDAE—LA  RUE 


221 


terior  to  cirrus-pouch.  Ovary  voluminous,  lobes  somewhat  winglike. 
Uterus  possessing  about  twenty  long  outpocketings  on  either  side.  Eggs 
provided  with  three  membranes.    Diameter  of  egg  0.024  mm. 

Habitat:     In  intestine   of  South  American  snakes  of  the  family 
Colubridae,  subfamily  Colubrinae. 


Host 

Locality 

Collector 

Authority 

Coluber  sp. 

Ophiomorphus  miliaris 
Ophiomorphus  miliaris 
Ophis  merremii 
Ophis  merremii 

Brazil 

Brazil* 

Brazil* 

Brazil* 

Brazil* 

Natterer 
Natterer 
Natterer 
Natterer 
Natterer 

Rudolphi,  1819:692 
Diesing,  1850:511 
Schwarz,  1908:28 
Diesing,  1850:511 
Schwarz,  1908:28 

This  species  was  first  described  by  Rudolphi  (1819 :692).  Since  this 
description  is  not  readily  accessible  to  many  workers  it  is  quoted  in  full : 

"Taenia  racemosa  R.  n,  sp.     Pone  Synops.  n.  52. 

"T.  Capite  obconico,  collo  brevi,  angustissimo,  articulis  planis,  elongatis,  fora- 
minibus  marginalibus  alternis  prominulis.  Hab.  In  intestinis  Colubri  n.  20.  speci- 
men sexpollicare  et  fragmenta  hujus  Taenia  Natterer  in  Brasilia  reperit, 

"Caput  antrorsum  dilatatum,  sive  obconicum,  osculis  orbicularibus,  aut  hemis- 
phaericis,  nam  satis  profunda  videntur,  anticis,  Collum  breve,  angustissimum. 
Articuli,  quos  vidi,  plani,  tenues,  elongati,  lineam  ad  sesquilineam  longi,  tertia 
lineae  parte  latiores,  foraminibus  marginalibus  alternis,  prominentibus. 

"Ovaria  singulorum  articulorum  lineam  mediam  fere  totam  sibi  vindicant,  ad 
cujus  latera  utrinque  maculae  exiguae  opacae  ita  digestae  sunt,  ut  ovaria  racemosa 
appareant. 

"Obs.  Caput  Taeniae  omphalodis  Synops.  n.  9.  articuli  vero  T.  tuberculatae  n. 
25.  ut  fragmenta,  quae  vidi,  Taeniam  illis  intermediam  reddant." 

Rudolphi  (1819:709)  established  the  name  Taenia  colubri  to  desig- 
nate a  few  proglottids  found  in  Coluber  sp.,  Brazil  by  Natterer.  His 
exact  words  are  here  quoted : 

"Taenia  Colubri.    Pone  Synops.  n.  140. 

Fragmenta  in  intestinis  Colubri  n.  12.  a  Natterero  in  Brasilia  reperta  Museo 
Viennensi  debeo  lineam  circiter  longa,  duodecim  ad  octodecim  articulis  constantia, 
latioribus  quam  longis,  subcuneatis,  angulis  plus  minus  exstantibus,  capite  destituta. 

"Utrum  vere  colubrina,  an  ex  ave  quadam  deglutita  forsan  residua?" 

Dujardin  (1845:610)  added  nothing  to  Rudolphi 's  description. 
Diesing  (1850:511)  gave  a  short  diagnosis  of  a  form  from  the  museum 
at  Vienna  which  he  identified  as  Taenia  racemosa  Rud.    His  description 

♦Material  found  in  Vienna  Museum. 


£22  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [222 

varies  so  widely  from  that  of  Rudolphi  and  his  list  of  hosts  covers  such 
a  range  of  families  that  his  words  are  quoted  in  full: 

"Taenia  racemosa  Rudolphi.  Caput  magnum  tetragonum,  acetabulis  angu- 
laribus  subterminalibus  v.  terminalibus,  subovatis  v.  cordatis.  Collutn  nullum. 
ArticuH  supremi  brevissimi,  subsequentes  longiores  et  latiores,  angulis  rotundatis, 
ultimi  longi  parallelopipedi  angustiores.  Penes  filiformes  basi  incrassati,  margi- 
nales  vage  alterni.     Longit.  2"  —  ij^';  latit.  med.  i  —  3"';  ultim.  1"'. 

"Taenia  racemosa  Rudolphi:  Synops.  App.  692 — Dujardin:  Hist.  nat.  des 
Helminth.  610. 

"Habitaculum.  Ophiomorphus  miliaris,  Martio.  —  Eunectes  Scytale,  Octobri, 
Februario  et  Junio.  —  Bothrops  Mararacca,  Martio  et  Aprili.  —  Ophis  Merremii, 
Januario,  in  Brasilia  {N alter er)  :  in  intestinis.  M.  C.  V." 

Von  Linstow  (1878:183)  did  no  more  than  refer  to  Rudolphi  (1819) 
and  Diesing  (1850).  Barrois  (1898:3)  was  convinced  that  Taenia  race- 
mosa Rud.  belonged  to  the  genus  Ichthyotaenia  for  he  had  examined  a 
head  and  some  ripe  proglottids  ''provenant  du  type  primitif  de  Rudol- 
phi." He  did  not  describe  the  species.  Liihe  (1898:652)  stated  that 
Taenia  racemosa  belonged  to  the  genus  Ichthyotaenia.  Marotel  (1899: 
34)  quoted  Diesing 's  (1850)  description  of  this  species  which  he  stated 
was  too  incomplete  to  permit  a  precise  determination.  Parona  (1901: 
3)  reported  some  specimens  collected  by  Dr.  Adolf  Lutz  from  Bothrops 
(Lachnis)  lanceolata,  Sao  Paulo,  Brazil  under  the  name  of  Ichthyotac' 
nia  racemosa  Rud.  These  specimens  which  have  been  examined  by  the 
writer  prove  to  belong  to  the  species  Ophiotaenia  calmettei  Barrois. 
Shipley  (1905:101)  reported  some  cestodes  from  Eunectes  murinus 
Wagler  as  Taenia  racemosa-  Rud.  Professor  H.  B.  "Ward  secured  some 
specimens  labelled  with  this  name  from  Mr.  Shipley.  Upon  examina- 
tion they  proved  to  be  specimens  of  Crepidobothrium  gerrardii  (Baird). 
Schwarz  (1908:28-29)  secured  some  material  from  Barrois  which  the 
latter  had  obtained  from  Dr.  E.  von  Marenzeller  of  Vienna.  He  also 
secured  additional  material  from  Dr.  E.  von  Marenzeller.  Upon  this 
material  he  based  his  description  of  Taenia  racemosa  Rud.  He  did  not 
definitely  state  the  name  of  the  host  from  which  his  material  was  col- 
lected but  evidently  quoted  Diesing 's  (1850)  list  of  hosts.  La  Rue 
(1911:481)  included  this  form  in  a  list  of  species  of  the  genus  Ophio- 
taenia. 

It  is  possible  that  T.  coluhri  Rud.  collected  by  Natterer  in  Brazil 
from  Coluber  sp.  is  a  sjTionym  of  T.  racemosa  Rud.  also  collected  by 
Natterer  in  Brazil  from  a  species  of  Coluber.  Rudolphi 's  description 
of  T.  coluhri  is  insufficient  to  place  the  species,  nor  is  there  any  data  as 
to  exact  host  and  locality  that  would  fix  it.    Diesing  (1850:558)  states 


223]  PROTEOCEPHALIDAE—LA  RUE  22Z 

that  T.  colubri  was  collected  by  Natterer  in  Brazil  from  Ophiomorphus 
poecilogyrus.  He  gave  no  descriptive  data  and  nothing  that  would  be 
of  assistance  in  placing  Rudolphi's  species.  Diesing's  (1850)  diagnosis 
of  Taenia  racemosa  Rud,  does  not  agree  with  Rudolphi's  description  of 
that  species.  His  statements  in  regard  to  the  tetragonal  head,  the  ter- 
minal or  subterminal  angular  suckers  which  are  subovate  or  cordate 
fits  the  diagnosis  of  Crepidobothrium  gerrardii  but  not  that  of  Ophio- 
taenia  racemosa.  The  latter  species  has  orbicular  suckers.  Likewise 
Diesing's  statements  that  there  is  no  neck,  that  the  first  proglottids  are 
very  short,  following  ones  larger  and  broader,  with  rounded  angles,  the 
last  proglottids  long  parallelopipeds  and  somewhat  narrow  agree  much 
better  with  the  descriptions  of  C.  gerrardii  than  with  that  of  0.  race- 
mosa. Diesing's  form  was  considerably  larger  than  Rudolphi's.  No 
completely  protruded  cirrus  was  noted  in  the  writer's  material  of  C. 
gerrardii  hence  the  cirri  of  the  two  forms  cannot  be  compared.  More- 
over, Diesing  lists  Eunectes  scytale  as  a  host  of  Taenia  racemosa.  Eunec- 
tes  scytale  is  a  synonym  of  Eunectes  murinus,  a  species  in  which  C. 
gerrardii,  but  no  other  Proteocephalid,  has  been  found.  It  seems  prob- 
able therefore  that  Diesing's  diagnosis  of  Taenia  racemosa  was  based  on 
the  material  from  Eunectes  murinus  (scytale). 

A  further  analysis  of  Diesing's  list  of  hosts  of  Taenia  racemosa 
shows  that  besides  the  Boidae,  represented  by  Eunectes  murinus,  the 
Colubridae  are  represented  by  two  species  and  the  Viperidae  by  one 
species.  From  this  evidence  it  seems  probable  that  Diesing's  Taenia 
racemosa  must  have  included  several  species,  for  among  the  species  of 
Ophiotaenia  from  snakes  which  have  been  adequately  described  in  recent 
years  no  species  has  been  found  in  hosts  belonging  to  different  families. 
A  list  of  these  species,  their  hosts,  and  their  distribution  is  found  else- 
where. Another  fact  to  be  remarked  about  these  Proteocephalid  species 
from  snakes  is  that  the  individuals  which  infest  the  Boidae  are  larger 
than  the  individuals  of  the  species  which  infest  the  Viperidae  and 
these  in  turn  are  larger  than  those  that  infest  the  Colubridae.  By  the 
term  size  the  writer  means  size  of  head,  size  of  suckers,  breadth  of  neck, 
breadth  and  length  of  proglottids  and  length  of  strobila.  In  certain 
of  these  characters  the  writer's  general  statement  fails  but,  considered 
in  a  broad  way,  it  is  true.  This  is  an  additional  reason  for  supposing 
that  Diesing  included  several  species  under  the  one  name  Taenia 
racemosa. 

Concerning  hosts  of  Taenia  racemosa  Schwarz  (1908:28)  says: 
"Als  Wirte  werden  angegeben:  Ophiomorphus  miliaris,  Ophis  Merre- 
mii,  Bothrops  jararacca  (for  mararacca)  und  Eunectes  scytale,  Schlan- 
gen,  die  in  Brazilien  vorkommen".    This  seems  to  be  but  a  restatement 


224  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [224 

of  Diesing's  list  of  hosts.  Nowhere  does  Sehwarz  state  definitely  the 
exact  host  from  which  his  specimens  were  taken.  A  study  of  his  de- 
scription in  comparison  with  other  species  from  snakes  shows  that  his 
specimens  are  most  closely  related  to  the  species  from  Colubrinae  yet 
they  are  distinct  from  any  of  those  species.  It  seems  then  quite  prob- 
able that  his  specimens  came  from  one  of  the  Colubrinae,  Coluber  sp. 
and  possibly  Ophiomorphus  miliaris  or  Ophis  merremii.  They  surely 
did  not  come  from  Eunectes,  and  it  is  highly  improbable  that  they  came 
from  Bothrops,  one  of  the  Viperidae.  The  emended  list  of  hosts  of 
Taenia  racemosa  then  contains  these  names,  Coluber  sp.,  Ophiomorphus 
miliaris  and  Ophis  merremii.  Further  questions  regarding  Sehwarz 's 
Taenia  racemosa  arise.  Is  his  species  identical  with  Eudolphi's  Taenia 
racemosa f  Could  Sehwarz  have  had  any  of  Rudolphi's  type  material? 
At  the  writer's  request  Professor  Ward  wrote  to  Professor  E.  von  Mar- 
enzeller  at  Vienna  for  information  concerning  the  specimens  of  Taenia 
racemosa  which  he  had  sent  to  Barrois  and  to  Sehwarz.  Professor  von 
Marenzeller  replied  that  all  of  Rudolphi's  types  were  in  the  Museum  at 
Berlin.  It  is  therefore  impossible  that  Sehwarz  or  "Barrois  had  Rudol- 
phi's type  specimens  for  examination  and  the  question  of  the  identity 
of  Sehwarz 's  Taenia  racemosa  and  Rudolphi's  species  of  that  name  re- 
mains open  and  must  so  remain  until  Rudolphi's  types  are  re-examined, 
if  they  still  exist.  Meanwhile  the  writer  assumes  that  the  Taenia  race- 
mosa of  Sehwarz  and  Rudolphi  are  identical. 

Sehwarz 's  material  all  came  originally  from  the  Museum  at  Vienna, 
some  directly  through  von  Marenzeller,  and  some  through  von  Maren- 
zeller to  Barrois,  then  from  Barrois  to  Sehwarz.  As  hosts  he  mentioned 
four  species  of  South  American  snakes  two  of  which  have  been  ruled 
out  in  the  above  discussion,  leaving  either  or  both  Ophis  merremii  and 
Ophiomorphus  miliaris  as  probable  hosts.  If  the  specimens  were  from 
Ophis  merremii  they  were  probably  collected  by  Natterer  in  Brazil. 

The  following  description  is  based  on  the  work  of  Sehwarz  (1908). 
The  observed  length  of  specimens  was  160  mm.,  length  of  ripe  proglottids 
2  mm.  and  the  breadth  of  the  same  1  mm.  The  scolex  (Fig.  140)  is 
readily  distinguished  from  the  strobila.  It  bears  four  nearly  circular 
suckers  which  measure  0.270-0.300  mm.  in  diameter.  The  scolex  has  a 
diameter  of  0.540  mm. 

The  sexual  organs  (Fig.  191)  agree  in  arrangement  with  the  genital 
organs  of  other  Ophiotaenia.  The  testes  are  of  exceptional  size,  0.072- 
0.078  mm.  in  diameter,  appearing  as  large  spheres.  They  are  not  limited 
to  the  side  fields  but  are  scattered  irregularly  through  the  whole  area 
of  the  segment.  The  testicular  field  is  not  interrupted  at  the  anterior 
or  posterior  margin  of  the  segment.    Testes  number  about  100-120.    The 


225]  PROTEOCEPHALIDAE—LA  RUE  225 

ductus  ejaculatorius  takes  a  sinuous  course  through  the  cirrus-pouch 
but  it  forms  no  coils.  The  cirrus-pouch  lies  perpendicular  to  the  longi- 
tudinal axis  of  the  proglottid  and  reaches  almost  to  the  median  line  of 
the  same. 

Female  organs  are  as  in  other  members  of  the  genus.  The  vagina 
opens  either  anterior  or  posterior  to  the  cirrus-sheath.  The  ov.ary  is 
voluminous,  bilobed,  joined  at  the  middle  by  a  mid-piece.  The  ovarian 
lobes  are  somewhat  plumper  in  form  than  are  those  of  0.  marenzelleri. 
The  uterus  forms  about  20  long  diverticula  on  either  side.  In  Schwarz's 
drawing  (reproduced  Fig.  191)  the  genital  pore  is  marginal  and  at  the 
end  of  the  first  third  of  the  proglottid.  The  eggs  are  round  and  have 
three  membranes.  No  booklets  are  to  be  found  on  the  outer  membrane. 
Schwarz  does  not  state  whether  the  diameter  of  0.024  mm.  is  the  diame- 
ter of  the  entire  egg  or  of  the  embryo. 

Ophiotaenia  racemosa  is  a  much  smaller  species  than  Crepidohoth- 
rium  gerrardii.  Its  head  and  suckers  are  smaller,  the  proglottids  shorter 
and  narrower.  The  form  of  the  suckers  is  greatly  different  in  the  two 
species.  0.  racemosa  differs  from  0.  calmettei  in  its  smaller  head, 
smaller  suckers,  in  the  distribution  of  testes  and  in  the  number  of  uterine 
pouches.  0.  racemosa  is  likewise  much  smaller  than  0.  marenzelleri  and 
O.  grandis  in  length,  breadth,  size  of  head,  and  size  of  suckers.  It  also 
has  fewer  testes.  0.  racemosa  differs  from  0.  timer esuri  in  being 
somewhat  smaller.  Its  chief  differences  lie  in  the  position  of  the  testes 
and  in  the  character  of  the  cirrus  and  vagina.  0.  racemosa  differs  from 
0.  pigmentata  in  having  larger  suckers.  0.  pigmentata  is  too  poorly 
described  to  furnish  more  diagnostic  differences.  0.  racemosa  differs 
from  0.  nattereri  and  0.  perspicua  in  having  a  larger  head,  larger  suck- 
ers,  a  different  arrangement  of  testes,  and  smaller  proglottids. 


226  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [226 


OPHIOTAENIA  PIGMENTATA  (von  Linstow) 

[Fig.  160] 

1908:    Ichthyotdenia  pigmentata      von  Linstow      1908:85 
1911:     Ophiotaenia  pigmentata         La  Rue  1911:481 

Von  Linstow  (1908)  described  this  species  from  a  specimen  which 
Edward  Jacobson  had  collected  in  Java  (Semarang).  La  Rue  (1911: 
481)  put  this  form  in  the  genus  Ophiotaenia.  Unfortunately  the  speci- 
men was  immature  so  von  Linstow 's  description  is  necessarily  incom- 
plete.   His  description  reads: 

"Ichthyotaenia  pigmentata,  n.  sp. — Aus  der  Schlange  Psammodynastes  pulver- 
ulentus  Fisch. — Darm. 

Lange  32mm.,  Breite  vom  0.51mm.,  hinten  1.78mm.;  ein  •sehr  lange  Strecke 
ist  ungegliedert,  nur  der  letzte  2.smm.  lange  Theil  zeigt  deutliche  Proglottiden- 
bildung;  es  ist  nur  ein  noch  voUig  entwickeltes  Exemplar  vorhanden.  Der  Scolex 
ist  abgerundet  und  nicht  breiter  als  die  folgende  Strecke;  die  Lange  des  Scolex 
betragt  0.57  mm. ;  die  kreisrunden  Saugnapfe  messen  o.i8mm. ;  ein  Rostellum  und 
Haken  fehlen.  Die  Glieder  am  Hinterende  sind  0.13mm.  lang;  ihre  Breite  betragt 
1. 1  mm.,  die  Dicke  0.25mm.  Die  Cuticula  ist  0.0052mm.  dick  und  tragt  an  ihrer 
Aussenseite  einen  0.0065mm.  dicken  Stabchenbesatz.  Die  Muskulatur  ist  stark 
entwickelt  und  man  unterscheidet  6  verschiedene  Muskelarten,  i,  Ring-und  2,  Langs- 
muskeln  in  diinnerer  Lage  unter  der  Cuticula ;  an  Parenchymmuskeln  3,  aussere,  4. 
innere"  Langsmuskeln,  letztere  bestehen  aus  Muskelbiindeln,  5.  Dorsoventral ;  und 
6.  Transversalmuskeln.  In  jedem  Querschnitt  erkannt  man  etwa  20  Langsgef asse ; 
die  Hauptlangensnerv-enstamme  verlaufen  1/5  des  Querdurchmessers  der  Proglot- 
tide  vom  Rande  entfemt;  massenhaft  finden  sich  dunkel  pigmentirte,  meistens 
eiformige,  durchschnittlich  0.026mm.  lange  und  0.021mm.  breite  Kalkkorperchen. 
Nur  in  den  letzten  Proglottiden  findet  man  eine  Anlage  der  Geschlechtsorgane, 
besonders  deutlich  ist  jederseits  nach  innen  vom  Nerv  der  rundliche  Dotter stock; 
trotz  der  geringen  Entwicklung  der  Geschlechtsorgane  ist  es  zweifellos  dass  die 
Art  zu  Ichthyotaenia  gehort". 

This  is  probably  a  species  of  Ophiotaenia.  The  presence  of  lateral 
vitellaria  removes  it  from  the  genus  Oochoristica.  It  must,  however,  be 
placed  in  the  list  of  incompletely  described  species.  .  ^ 


227]  PROTEOCEPHALIDAE—LA  RUE  227 


1898 
1898 
1908 
1911 


OPHIOTAENIA  TRIMERESURI  (Parona) 
[Figs.  106-108,  141,  142,  192,  193] 

Taenia  trimeresuri  Parona  1898:7-11 

Ichthyotaenia  trimeresuri  Liihie  1898:652 

Ichthyotaenia  trimeresuri  Schwarz  1908:33-35 

Ophiotaenia  trimeresuri  La  Rue  1911:481 


Specific  Diagnosis:  Cliaracters  of  genus.  Length  up  to  105  mm. 
Iklaximum  breadth  0.75-1.5  mm.  Scolex  unarmed,  without  rostellum, 
broader  than  necli,  breadth  0.75  mm.  Suckers  prominent,  hemispherical, 
muscular,  0.16-0.25  mm.  in  diameter,  situated  anteriorly.  No  fifth 
suclier.  Necli  short,  three  times  length  of  head,  breadth  0.25-0.5  mm. 
First  proglottids  broader  than  long.  IVIature  proglottids  quadrate  or 
longer  than  broad.  Angles  of  proglottids  not  prominent.  Strobilation 
indistinct. 

Genital  pore  situated  near  middle  of  proglottid  margin,  irregularly 
alternating  in  position.  No  genital  papilla.  Genital  sinus  if  present 
very  shallow.  Testes  100-108  in  number,  measuring  0.063  by  0.027  to 
0.080  by  0.027  mm.,  situated  in  two  fields  not  near  vitellaria.  All  testes 
anterior  to  ovary.  Mass  of  coils  of  vas  deferens  not  large.  Ductus 
ejaculatorius  much  coiled.  Cirrus  muscular.  Many  coils  of  ductus 
ejaculatorius  in  base  of  protruded  cirrus.  Cirrus-pouch  about  0.270- 
0.330  mm.  long  by  0.136  mm.  broad.  Ratio  of  length  of  cirrus-pouch 
to  proglottid  breadth  1 :4  or  2 :5.  Vagina  anterior  or  posterior  to  cirrus- 
pouch,  not  crossing  same.  Lumen  of  vagina  variable  in  size.  Sphincter 
vaginae  present.  Lobes  of  ovary  flattened,  elongated,  narrow.  Vitel- 
line follicles  small.  Uterus  when  fully  developed  with  20-30 (?)  lateral 
pouches.    No  ripe  eggs  observed. 

Habitat :  In  the  intestine  of  Trimeresurus  formosus  (type  host)  ; 
Island  of  Mentawei,  East  Indies  (type  locality). 

Type:  Specimens  in  Professor  C.  Parona 's  collection  labelled 
" Arynchotaenia  trimeresuri  Par.,  Trimeresurus  formosus,  (Mentawei)." 
Also  twb  toto  preparations  of  the  same  material  in  the  collection  of 
Professor  H.  B,  Ward. 

Parona  ( 1898 :7-ll )  first  described  and  figured  this  species  without 
discovering  its    relation    to    the    Proteocephalidae.     Liihe    (1898:652) 


228  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [228 

stated  that  this  species  belonged  to  the  Ichthyotaenia.  In  a  footnote  he 
says,  "Herr  Prof.  Parona  hat  mir  die  Original-exemplare  der  Art  in 
uneigenniitzigster  Weise  zur  VerfUgung  gestellt,  so  dass  ieh  die  Zuge- 
horigkeit  derselben  zu  der  Gattung  Ichthyotaenia  feststellen  konnte." 
Sehwarz  (1908:33-35)  had  no  specimens  of  this  species  for  examination. 
He  rewrote  Parona 's  description  but  was  able  to  add  nothing  new.  He 
believed  this  form  to  be  a  species  of  Proteocephalus  and  called  attention 
to  the  fact  that  the  lower  part  of  the  uterus  in  Parona 's  drawing  was 
probably  the  ovary  and  that  the  testes  would  be  found  in  the  anterior 
region  of  the  proglottid  where  they  were  frequently  difficult  to  see. 
La  Rue  (1911:474)  stated  that  this  species  belonged  with  the  genus 
Ophiotaenia  and  gave  a  few  of  its  characters. 

Thanks  to  Professor  H.  B.  Ward  who  secured  this  material  from 
Professor  Parona  for  the  purpose  of  this  study  the  writer  has  been  able 
to  give  this  species  a  more  careful  description  than  has  hitherto  been 
attempted.  The  material  consists  of  about  a  half  dozen  pieces  among 
which  no  head  could  be  found.  This  material  is  labelled  "  Arynchotae- 
nia  trimeresuri  Par.,  Trimeresurus  formosus  (Mentawei) ".  From  it 
two  toto  preparations  were  made.  These  together  with  the  alcoholics 
and  Parona 's  original  description  form  the  basis  of  the  following 
description. 

In  the  material  at  the  writer's  disposal  the  head,  neck  and  first 
proglottids  were  missing.  The  pieces  measured  10,  20,  60  and  70  mm. 
long  by  a  maximum  breadth  of  about  1.0  mm.  Parona  in  a  table  gave 
data  concerning  the  length  and  breadth  of  five  specimens  with  heads 
and  five  without.  The  length  of  those  with  heads  ranged  from  12  to  50 
mm.  and  the  maximum  breadth  from  0.75  to  0.1  mm.  while  the  headless 
specimens  measured  33-105  mm.  in  length  by  a  maximum  breadth  of 
0.75-1.5  mm.  The  scolex  is  spheroidal,  has  neither  hooks  nor  rostellum, 
and  is  broader  than  the  neck.  Its  diameter  is  about  0.75  mm.  The 
suckers  are  prominent  and  are  situated  in  the  anterior  part  of  the  head. 
They  are  hemispherical,  strongly  muscular,  and  have  a  diameter  of  0.16- 
0.25  mm.  The  aperture  of  the  sucker  is  0.11-0.16  mm.  Parona 's  draw- 
ings of  the  scolex  are  reproduced  (Figs.  141,  142). 

The  neck  is  about  three  times  as  long  as  the  scolex.  Its  breadth 
varies  from  0.25  to  0.5  mm.  The  first  proglottids  tho  broader  than  long 
are  not  very  short.  These  become  successively  longer,  the  last  being  long- 
est. The  increase  in  length,  however,  is  not  constant  because  short  pro- 
glottids are  to  be  found  between  long  ones.  Nor  is  the  breadth  of  the 
strobila  constant.  Some  proglottids  are  about  quadrate  while  others  are 
2-3-4  times  longer  than  broad.  Proglottid  limits  are  poorly  defined  ren- 
dering the  strobilation  indistinct.    The  angles  of  the  proglottids  are  not 


229]  PROTEOCEPHALIDAE—LA  RUE  229 

prominent  hence  the  worm  has  the  appearance  of  a  continuous  ribbon. 
The  description  thus  far  is  based  on  the  original  description  as  given  by 
Parona  (1898:8-9).  In  the  toto  preparations  which  the  writer  has 
studied  there  were  a  few  mature  proglottids  one  of  which  is  delineated 
(Fig.  108).  The  uterus  of  this  proglottid  contained  a  few  eggs  but 
these  have  been  omitted  in  the  drawing.  This  segment  measured  1.7 
mm.  long  by  0.9  mm.  broad.  It  was  thin  and  flat  but  considerably 
thicker  than  proglottids  of  an  equal  state  of  development  from  0.  per- 
spicua  or  0.  natterer. 

A  genital  pore  is  situated  near  the  middle  of  the  margin  of  each 
proglottid.  It  alternates  irregularly  from  left  to  right.  There  is  no 
genital  papilla  nor  is  the  pore  marked  by  a  deep  depression  as  is  some- 
times the  case  in  0.  grandis.  The  vagina  and  cirrus-pouch  open  very 
near  each  other  but  in  this  toto  preparation  it  was  impossible  to  tell 
whether  there  was  a  common  genital  sinus.  If  present  at  all  it  was  very 
shallow.  The  other  drawings  (Figs.  106,  107)  do  not  show  such  a 
structure. 

Testes  (Fig.  108)  are  100-108  in  number.  Their  dimensions  are 
0.063  by  0.027  mm.  to  0.080  by  0.027  mm.,  the  long  axis  being  perpen- 
dicular to  the  long  axis  of  the  worm.  They  are  arranged  in  narrow 
bands  situated  well  away  from  the  vitellaria.  In  this  respect  the  species 
resembles  0.  calmettei.  None  of  the  testes  are  posterior  to  the  ovaries 
though  Parona 's  drawing  which  has  been  reproduced  (Fig.  193)  for 
purposes  of  comparison  shows  them  there.  The  vas  deferens  (Fig.  108) 
forms  a  small  mass  of  coils  in  the  mid-region  of  the  proglottid.  Within 
the  cirrus-pouch  is  the  much  coiled  ductus  ejaculatorius  (Fig.  107). 
The  ductus  passes  over  into  the  thicker- walled  and  more  muscular  cirrus. 
When  under  certain  conditions  the  cirrus  is  protruded  the  greater  part 
of  the  ductus  ejaculatorius  is  crowded  out  into  the  dilated  basal  part 
of  the  cirrus  (Figs.  106,  107).  In  this  condition  the  cirrus  is  very 
similar  to  that  described  and  figured  by  Schwarz  (1908)  for  0.  maren- 
zelleri.  He,  however,  claimed  that  the  whole  cirrus-pouch  was  evagi- 
nated.  In  this  he  misinterpreted  the  facts  for  only  the  cirrus  and  a 
part  of  the  ductus  are  pushed  out.  Schwarz 's  drawing  (reproduced  Fig. 
199)  shows  the  cirrus-pouch  in  its  normal  position.  In  0.  marenzelleri 
the  distal  half  of  the  evaginated  cirrus  is  filiform.  This  condition  has 
not  been  seen  in  the  present  species  yet  it  seems  highly  probable  that  a 
part  of  the  coils  of  the  ductus  can  be  pushed  through  the  basal  part  of 
the  cirrus  and  thus  form  a  filiform  cirrus.  Unless  this  be  the  case  it  is 
difficult  to  understand  how  copulation  can  be  possible.  The  large 
number  of  coils  of  ductus  ejaculatorius  in  this  species  furnishes  a  dif- 
ferentiating character  between  this  species  and  0.  marenzelleri.     The 


230  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [230 

cirms-poueh  (Fig.  107)  is  large  and  very  muscular.  Its  length  varies 
from  0.27  to  0.34  mm.  Its  breadth  is  about  0.136  mm.  and  its  length, 
according  to  Parona,  is  0.33  mm.  The  ratio  of  the  length  of  the  cirrus- 
pouch  to  the  breadth  of  proglottid  is  1 :4  or  2 :5. 

The  vagina  lies  either  anterior  or  posterior  to  the  cirrus-pouch,  but 
it  does  not  cross  the  latter.  Its  course  after  reaching  the  median  field 
is  frequently  quite  sinuous.  Near  its  opening  (Figs.  106,  107)  it  has  a 
very  heavy  sphincter  muscle.  In  some  parts  of  the  vagina  its  lumen  is 
narrow  while  in  other  regions  it  may  be  widely  dilated.  The  lobes  of 
the  ovary  are  flattened.  They  are  more  elongate  and  narrower  than 
in  0.  marenzelleri.  The  paired  vitelline  ducts  cross  the  ovary  on  the 
ventral  side.  The  uterus  in  mature  proglottids  is  a  median  tube  ex- 
tending the  length  of  the  proglottid.  No  ripe  proglottids  could  be 
examined  so  the  number  of  lateral  uterine  out-pocketings  could  not  be 
accurately  determined.  The  drawing,  however,  (Fig.  108)  indicates 
that  probably  about  20-30  lateral  pouches  would  be  developed  on  either 
side.    No  ripe  eggs  were  present  in  the  material  examined. 

The  description  and  drawing  of  the  mature  proglottid  shows  this 
to  be  a  species  which  falls  readily  into  the  group  of  Proteocephalids 
infesting  snakes.  The  writer's  description  and  drawings  do  not  agree 
very  well  with  Parona 's  except  as  to  size,  position  of  genital  pore  and 
length  and  shape  of  cirrus-pouch.  From  his  drawings  which  have  been 
reproduced  (Figs.  192,  193)  it  appears  that  he  has  confused  certain 
organs.  The  lower  portion  of  the  uterus  in  his  drawing  is  undoubtedly 
the  bilobed  ovary.  The  position  of  the  testes  posterior  to  the  ovary  in 
his  figure  cannot  be  explained  except  on  the  supposition  that  he  mistook 
parts  of  the  organs  af  the  interovarial  space  for  testes.  The  sinuous 
duct  which  he  shows  as  the  vas  deferens  extending  back  from  the  cirrus- 
pouch  to  the  ovary  is  not  the  vas  deferens  but  the  vagina  which  in  fact 
passes  below  the  coils  of  vas  deferens  before  reaching  the  middle  of  the 
proglottid.  The  mass  of  coils  of  the  vas  deferens  is  poorly  shown  at 
the  inner  end  of  the  cirrus-pouch.  The  opening  of  the  vagina,  the 
vaginal  sphincter  and  the  first  part  of  vagina  along  the  length  of  the 
cirrus-pouch  he  has  omitted  altogether  or  he  has  confused  these  with  a 
part  of  the  cirrus-pouch.  The  cirrus-pouch  as  he  drew  it  seems  to  be 
made  up  of  lamellae.  In  a  poor  preparation  the  cirrus-pouch  might 
have  had  this  appearance.  The  cirrus  within  the  cirrus-pouch  is  not 
properly  delineated.  The  uterus  in  an  early  stage  may  possibly  have 
had  the  single  anterior  lateral  pouch  on  either  side  as  he  has  shown  it. 
The  writer  has  not  seen  a  proglottid  as  nearly  ripe  as  this  one  was  and 
so  is  not  able  to  judge  as  to  the  actual  structure  of  the  uterus.  The 
testes  and  vitellaria  he  (Parona)  has  omitted  from  his  drawings.     It 


231] 


PROTEOCEPHALIDAE—LA  RUE 


231 


seems  highly  probable  that  Parona's  drawings  are  intended  to  represent 
the  same  species  as  do  the  writer's.  The  apparent  difference  probably 
came  about  through  Parona's  using  poor  preparations  for  study. 

0.  trimeresuri  (Parona)  in  size  is  much  smaller  than  0.  grandis, 
0.  marenzelleri  and  Crepidohothrium  gerrardii.  Moreover  in  number 
and  arrangement  of  testes,  size  and  proportions  of  cirrus-pouch  and 
cirrus  these  three  species  differ  greatly  from  0.  trimeresuri.  In  size 
this  species  is  more  nearly  related  to  0.  calmettei  but  in  the  latter 
species  the  relations  of  the  cirrus,  cirrus-pouch  and  vagina  are  different. 
The  testes  in  0.  trimeresuri  are  arranged  much  as  they  are  in  0.  cal- 
mettei but  the  size  of  the  head  and  the  suckers  differ  greatly.  0.  nat- 
tereri  and  0.  perspicua  are  smaller,  more  delicate  and  have  very  differ- 
ent relations  of  cirrus,  cirrus-pouch,  vagina,  and  testes.  This  form 
differs  from  any  other  species  thus  far  described  from  snakes  yet  it  has 
its  closest  affinities  with  0.  calmettei,  likewise  parasitic  in  one  of  the 
Crotalinae. 


1898 
1898 
1898 
1899 
1901 
1908 
1911 


OPHIOTAENIA  CALMETTEI  (Barrois) 
[Figs.  11,  109,  110,  155,  156,  197,  198] 

Ichthyotaenia  calmettei 
Ichthyotaenia  raillieti 
Ichthyotaenia  calmettei 
Ichthyotaenia  calmettei 
Ichthyotaenia  racemosa 
Ichthyotaenia  calmettei 
Ophiotaenia  calmettei 


Barrois 

1898:1-3 

Marotel 

1898 :99-101 

Liihe 

1898 :652 

Marotel 

1899 :34-42 

Parona 

1901 :3 

Schwarz 

1908 :24-26 

La  Rue 

1911 :481 

Specific  Diagnosis:  Characters  of  genus.  Observed  length  27-40, 
and  even  up  to  80  cm.  Maximum  breadth  0.97-1.2-2.0  mm.  Proglottids 
numerous,  first  broader  than  long,  subsequent  ones  quadrate  or  even 
much  longer  than  broad.  Mature  proglottids  about  0.85  ram.  square. 
Ripe  proglottids  2-3-4  mm.  long  by  1.0-1.2  mm.  broad.  Strobilation  not 
clear.  Scolex  spheroidal  or  tetragonal,  without  rostellum,  without  hooks, 
without  fifth  sucker.  Anterior  face  of  same  flattened,  with  small  eleva- 
tion at  center.  Head  1.0-1.3  mm.  broad,  0.60  mm.  long,  0.935  mm.  thick. 
Suckers  four,  somewhat  globular,  with  deep  cavity.  Diameter  of  sucker 
opening  0.12-0.17-0.185  mm.  Diameter  of  sucker  0.27-0.300-0.408  mm. 
Neck  0.580-0.980  mm.  broad  by  4-5-8  mm.  long. 

Genital  organs  typical  of  genus.  Genital  aperture  irregularly  alter- 
nating.   Situated  at  or  near  middle  of  proglottid  margin.    Genital  sinus 


ut 


ILLINOIS    BIOLOGICAL    MONOGRAPHS 


[232 


shallow.  Vagina  anterior  or  posterior  to  cirrus-pouch.  Vaginal  opening 
frequently  dorsal  to  cirrus-pouch.  Testes  130-160  in  number,  arranged 
in  two  lateral  fields.  Size  of  testes  0.026-0.04-0.06  by  0.053-0.07  mm. 
Vas  deferens  a  mass  of  coils  extending  from  cirrus-pouch  to  mid-field. 
Cirrus-pouch  0.25-0.29-0.33  mm.  long  by  0.100-0.136  mm.  broad.  Ratio 
of  length  of  cirrus  to  proglottid  breadth  1 :6-l  :5  in  mature  and  1 :4  in 
ripe  proglottids.  Ductus  ejaculatorius  with  numerous  coils.  Cirrus 
broad,  muscular,  not  seen  protruded. 

Vagina  broad  at  distal  end,  never  crossing  cirrus-pouch  in  course 
to  mid-field.  Course  of  vagina  to  interovarial  space  sinuous.  Sphincter 
yaginae  present.  Receptaculum  seminis  not  seen.  Ovarian  lobes  slender 
in  young  proglottids,  broad  in  old.  Vitellaria  with  small  follicles. 
Uterus  when  fully  developed  possessing  24-35  lateral  pouches  on  either 
side.  Uterine  pores  ventral,  2-4-5  in  number.  Eggs  provided  with 
three  membranes.  Outer  membrane  variable  in  size,  second  one  thicker, 
0.022-0.024  mm.  in  diameter,  inner  one  granular,  irregular,  0.014  mm., 
embryo  0.012  mm. 

Habitat:    In  intestine  of  Lachesis  (Bothrops)  lanceolatus  L. 


Host 


Lachesis  (Bothrops)  lanceolatus  L. 
Lachesis  {Bothrops)  lanceolatus  L. 
Lachesis  (Bothrops)  lanceolatus  L. 


Locality 

Collector 

Authority 

Martinique 

Calmette 

Barrois 

Martinique 

Guerin 

Marotel 

Sao  Paulo, 

Adolf  Lutz 

La  Rue   (the 

Brazil 

present  paper) 

Barrois'  (1898)  description  of  this  species  while  meager  and  un- 
accompanied by  drawings  agrees  fairly  well  with  the  descriptions  of 
later  investigators.  His  specimens  came  from  Lachesis  (Bothrops)  lan- 
ceolatus L.  from  Martinique  from  which  host  they  were  secured  by 
Professor  Calmette.  Later  in  the  same  year  Marotel  (1898)  briefly 
described  this  species  under  the  name  /.  raillieti.  His  specimens  came 
from  Lachesis  (Bothrops)  lanceolatus  L.  from  Martinique.  Liihe  (1898) 
called  attention  to  the  fact  that  I.  raillieti  was  a  synonym  of  /.  calmet- 
tei,  and  he  included  the  form  in  his  list  of  species  of  Ichthyotaenia. 
Marotel  (1899:34-42)  in  a  more  extended  paper  described  this  species. 
His  paper  is  accompanied  by  several  drawings. 

Parona  (1901)  listed  this  species  in  a  record  of  parasites  from 
South  America  under  the  name  of  Ichthyotaenia  racemosa.  His  speci- 
mens are  from  B.  lanceolatus  L.  from  Sao  Paulo,  Brazil.  Schwarz 
(1908:24-26)  added  but  little  to  the  work  of  Barrois  and  Marotel.    He 


233]  PROTEOCEPHALIDAE—LA  RUE  233 

determined  that  there  was  no  rostellum.  La  Rue  (1911:481)  included 
this  form  in  a  list  of  species  of  Ophiotaenia. 

This  study  is  based  on  material  which  Professor  H.  B.  Ward 
secured  for  the  writer's  study  from  Professor  Parona.  Professor 
Parona's  material  bore  the  label,  "T.  (Oochoristica)  racemosa  S.  Paulo, 
race.  A.  Lutz."  This  is  evidently  the  material  on  which  Parona  (1901) 
based  his  report.  There  he  stated  that  it  came  from  Bothrops  lanceo- 
latus  L.  The  material  consisted  of  seven  or  eight  pieces  to  one  of  which 
a  head  was  attached.  The  head  was  cleared  in  glycerine  and  studied 
in  that  condition  while  some  of  the  smaller  pieces  were  stained  and 
mounted  as  toto  preparations.  The  identification  of  the  material  was 
made  from  these  preparations.  The  report  is  also  based  upon  data 
secured  from  the  papers  of  Barrois  (1898),  Marotel  (1899),  and 
Schwarz  (1908). 

The  longest  piece  which  the  writer  observed  measured  270  mm.  long 
by  2.0  mm.  broad.  Barrois  (1898)  states  that  he  observed  a  length  of 
35-80  cm.  by  a  breadth  of  0.97  mm.  for  this  species.  Marotel's  (1899) 
specimens  measured  up  to  35-40  cm.  long  by  a  maximum  breadth  of  1.2 
mm.  Schwarz  (1908)  reported  a  breadth  of  1.5  mm.  The  strobila  is 
made  up  of  many  proglottids,  according  to  Barrois  as  many  as  289. 
The  proglottids  are  closely  joined  to  one  another.  As  a  rule  no  inter- 
segmental furrows  can  be  seen  with  the  unaided  eye.  In  a  single  piece 
the  strobila  was  strongly  contracted  and  here  the  strobilation  was  evi- 
dent. The  surface  of  many  of  the  proglottids  is  thrown  into  numerous 
longitudinal  folds. 

The  scolex  is  club-shaped  according  to  Barrois  and  Marotel,  sphe- 
roidal according  to  Schwarz.  The  figures  of  the  head  as  delineated  by 
Marotel  and  Schwarz  are  reproduced  (Figs.  155,  156).  Barrois  and 
Schwarz  reported  that  there  was  no  fifth  sucker  and  no  rostellum  while 
Marotel  thought  that  the  slight  elevation  at  the  middle  of  the  head 
(Fig.  156)  was  a  rudimentary  rostellum.  He  saw  no  fifth  sucker. 
Marotel  states  that  the  globular  suckers  are  directed  anteriad.  The 
head  (Fig.  11)  which  the  writer  observed  was  cleared  in  glycerine.  It 
presents  a  somewhat  tetragonal  face  with  a  small  elevation  at  its  center 
which  is  not  a  rostellum  and  upon  which  there  is  no  fifth  sucker.  The 
suckers  are  placed  in  the  corners  of  the  anterior  face  and  they  are 
directed  anteriad.  Slight  grooves  or  wrinkles  which  do  not  extend  to 
the  apex  partially  divide  the  head  into  quadrants.  The  suckers  are 
nearly  round  in  outline  and  they  lack  any  trace  of  the  inturned  lower 
margin  which  is  a  characteristic  of  the  suckers  of  Crepidohothrium 
gerrardii.  Suckers  measure  0.39-0.408  mm.  in  diameter  while  the  open- 
ings of  the  suckers  measure  0.17-0.185  mm.  in  diameter.     The  sucker 


234 


ILLINOIS    BIOLOGICAL    MONOGRAPHS 


[234 


2avity  is  deep.  This  head  is  1.19  mm.  broad,  0.60  mm.  long,  0.935  mm. 
thick.  The  comparative  table  which  follows  presents  the  data  on  the 
head  as  secured  by  Barrois,  Marotel,  Schwarz,  and  the  writer. 


Head 

Suckers 

Diameter 

of  sucker 

opening 

Authority 

Shape 

Size 

Shape 

Diameter 

Club-shaped 

breadth  1-1.3  mm. 
breadth  1.05  mm. 
breadth  i.omm. 
breadth  1.19  mm. 
length  0.60  mm. 
thickness  0.935  r"in- 

lo/iZ-ci.TA  mm 

Barrois  (1898:2) 

Club-shaped 
Spheroidal 
Somewhat  tet- 
ragonal 

globular 

0.27-0.30  mm. 
0.30-0.34  mm. 
0.39-0.408  mm. 

0.120  mm. 

0.120  mm. 

0.1 7-0. 185 

mm. 

Marotel  (1899:36) 
Schwarz  (1908:24) 

round  in 
outline 

La  Rue  (the 
present  paper) 

The  neck  of  the  specimen  observed  by  the  writer  measured  about 
5-8  mm.  long  by  0.980  mm.  broad.  Its  limits  could  not  readily  be  de- 
termined in  a  glycerine  preparation.  Marotel  states  that  the  neck  is  4 
or  5  mm.  long  and  0.580  mm.  broad  at  its  narrowest  place.  Barrois  and 
Schwarz  do  not  state  its  dimensions.  The  first  proglottids  are  much 
broader  than  -long.  These  increase  in  length  more  rapidly  than  in 
breadth.  Marotel  (1899)  states  that  at  20  mm.  from  the  head  the  pro- 
glottids measure  0.255  mm.  long  by  0.63  mm.  broad ;  at  25  mm.  0,65  by 
0.65  mm.  i.  e.,  they  are  quadrate ;  at  30  mm.,  0.71  mm.  long,  0.67  mm. 
broad ;  at  40  mm.,  1.45  by  0.75  mm.,  and  the  last  proglottids  are  3  to  4 
mm.  long  by  1.0  mm.  broad.  In  material  examined  by  the  writer  nearly 
mature  proglottids  measure  as  much  as  0.9  mm.  broad  by  0.55  mm. 
long  and  mature  proglottids  about  0.85  mm.  square  or  some  of  them 
may  be  a  little  longer  than  broad.  Ripe  proglottids  measure  2-3  mm. 
long  by  1.0-1.2  mm.  in  breadth.  In  some  contracted  regions  of  the 
strobila  the  breadth  may  measure  as  much  as  2.0  mm. 

The  sexual  aperture  is  irregularly  alternating,  situated  in  mature 
proglottids  at  or  slightly  anterior  to  the  middle  of  the  margin.  In  ripe 
proglottids  it  may  be  either  anterior  or  posterior  to  the  middle.  Both 
cirrus  and  vagina  open  into  a  common  genital  sinus  which  is  very 
shallow.  The  vagina  lies  anterior  or  posterior  to  the  cirrus-pouch  with 
almost  equal  frequency.  In  ripe  proglottids  frequently  and  in  mature 
proglottids  more  rarely,  the  vagina  may  open  dorsal  to  the  cirrus- 
pouch. 

The  testes  (Figs.  109,  110)  number  from  130  to  160  in  the  writer's 
preparation  and  these  numbers  are  also  about  the  limits  observed  by 
the  other  investigators.  They  measure  from  0.026  to  0.037  mm.  by  0.053 
to  0.063  mm.  in  the  writer's  preparations.  Others  report  them  as  measur- 


235]  PROTEOCEPHALIDAE—LA  RUE  235 

ing  0.06  by  0.04  mm.,  0.062-0.072  mm.,  and  0.07  by  0.04  mm.  They  are 
arranged  in  two  fields  which  in  elongated  proglottids  are  near  the 
ventral  excretory  vessels  while  in  contracted  proglottids  (Figs.  109, 
110)  they  are  some  distance  from  the  ventral  vessels.  In  the  former 
ease  the  testes  of  each  field  are  arranged  in  two  irregular  rows  while 
in  the  latter  case  the  testes  of  each  field  are  irregularly  arranged  in  a 
broad  zone.  The  testes  are  dorsal  to  the  uterus.  The  vas  deferens 
forms  a  mass  of  coils  which  extend  from  the  cirrus-sheath  nearly  to  the 
middle  of  the  proglottid.  The  writer's  drawings  (Figs.  109,  110)  show 
it  more  heavily  developed  than  do  the  figures  of  Marotel  (1899)  which 
are  reproduced  (Figs.  197,  198). 

The  cirrus-pouch  is  relatively  short  and  broad,  0.25-0.29  mm.  long 
by  0.100-0.136  mm.  broad.  In  the  broad  mature  and  ripe  proglottids 
it  extends  but  a  short  distance  within  the  vitellaria.  The  ratio  of  its 
length  to  the  breadth  in  mature  proglottids  is  about  1:6,  in  ripe  pro- 
glottids about  1 :5  or  1 :5.5,  in  the  ripest  proglottid  observed  about  1 :4. 
Schwarz  states  that  the  cirrus-pouch  is  about  0.33  mm.  long.  The 
ductus  ejaculatorius  (Fig.  109)  has  numerous  coils.  The  unprotruded 
cirrus  is  broad  and  muscular.  The  writer  has  not  seen  it  protruded 
and  no  other  investigator  reports  having  seen  it  thus.  An  examination 
of  Marotel's  figures  which  have  been  reproduced  (Figs.  197,  198)  shows 
fewer  coils  of  ductus  ejaculatorius  and  of  vas  deferens  than  do  the 
drawings  of  the  writer  (Figs.  109,  110).  This  can  be  explained  by  the 
assumption  that  Marotel  could  not  follow  these  structures  out  com- 
pletely in  his  preparation  which  he  states  was  made  from  material  in 
a  poor  state  of  preservation.  The  material  upon  which  the  writer 
worked  was  in  a  fair  state  of  preservation  and  in  preparations  made 
from  it  these  ducts  could  be  traced  with  ease. 

The  vagina  at  its  distal  end  is  broad,  measuring  0.09-0.12  mm.,  a 
breadth  due  in  part  to  the  breadth  of  its  lumen  and  in  part  to  the 
thickness  of  its  sphincter  vaginae.  In  this  region  the  vagina  is  nearly 
as  broad  as  is  the  cirrus  in  its  basal  portion.  In  its  course  to  the  middle 
of  the  proglottid  the  vagina  never  crosses  the  cirrus-pouch.  Arrived 
at  the  middle  of  the  segment  it  bends  sharply  and  takes  a  sinuous 
course  posteriad  to  the  interovarial  space  which  it  enters  after  passing 
over  the  dorsal  side  of  the  ovary,  not  the  ventral  as  Marotel  figures  it. 
There  are  no  coils  of  the  vagina  anterior  to  the  ovary.  A  receptaculum 
seminis  has  not  been  observed  tho  from  its  occurrence  in  other  species 
in  the  genus  its  presence  in  this  species  may  be  inferred.  The  lobes 
of  the  ovary  in  nearly  mature  proglottids  are  more  slender  and  more 
pointed  at  the  extremities  than  in  mature  and  ripe  proglottids.    In  well 


236  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [236 

elongated  segments  the  lobes  of  the  ovary  assume  the  form  figured  by 
Marotel  (Figs.  197,  198). 

The  vitellaria  are  lateral,  and  made  up  of  small  follicles.  The 
vitelline  ducts  cross  the  ventral  surface  of  the  ovary  instead  of  lying 
some  distance  anterior  to  it  as  Marotel  figured  them.  In  mature  pro- 
glottids  the  uterus  is  a  median  tube.  The  early  development  of  the 
lateral  pouches  (Figs.  110,  198)  may  be  traced  as  small  somewhat 
excentric  bulbous  enlargements  of  the  median  tube.  This  is  not  essen- 
tially different  from  0.  filaroides  and  0.  perspicua.  These  enlarge- 
ments may  attain  some  size  before  any  uterine  eggs  appear.  In  early 
stages  there  is  considerable  resemblance  to  the  condition  shown  in  Mar- 
otel's  figure  (reproduced  Fig.  198)  of  the  ovigerous  segment  of  0. 
calmettei.  Later,  how^ever,  many  of  these  pouches  (Fig.  109)  extend 
toward  the  sides  and  the  uterus  resembles  that  of  other  species  of  the 
genus.  The  walls  of  these  pouches  are  not  very  easily  observed  so 
Marotel  may  have  overlooked  them  in  his  specimens.  The  pouches 
number  24-35  on  either  side.  Two  to  four  or  five  ventral  uterine  pores 
were  observed  in  some  of  the  riper  proglottids.  The  uterine  eggs  in 
the  alcoholic  material  are  surrounded  by  three  membranes.  The  outer- 
most very  thin  hyaline  membrane  measures  from  0.024  to  0.026  mm.  The 
second,  a  thick  and  homogeneous  membrane,  measures  0.022  mm.  The 
inner  membrane  is  thick,  granular  and  more  or  less  irregular  in  outline. 
It  measures  0.014  mm.  while  the  six-hooked  embryo  measures  0.012  mm. 
in  diameter.  Marotel  (1899)  states  that  the  eggs  are  globular,  with 
two  membranes,  the  outermost  one  being  thin  and  membranous,  0.065 
mm.  in  diameter,  and  the  ,other,  a  homogeneous  and  somewhat  thick 
membrane,  0.024  mm.  in  diameter.  This  membrane  corresponds  in  size 
and  description  to  the  one  which  the  writer  calls  the  middle  membrane- 
He  further  states  that  the  embryo  is  granular  but  does  not  give  its 
dimensions. 

This  species  varies  from  the  0.  racemosa  described  by  Schwarz 
(1908)  in  the  much  larger  size  of  the  head,  the  larger  size  of  the  suck- 
ers, and  the  relative  prominence  of  the  same.  The  number  and  arrange- 
ment of  the  testes  are  radically  different.  In  the  length  of  the  cirrus- 
pouch,  in  the  number  of  coils  of  ductus  ejaculatorius  they  differ  greatly. 
In  the  size  of  the  distal  end  of  the  vagina  there  is  considerable  differ- 
ence. The  character  of  the  diverticula  of  the  uterus  is  not  the  same. 
In  this  species  the  uterus  extends  back  to  the  ovary  while  in  0.  race- 
mosa as  described  by  Schwarz  it  does  not.  The  vitellaria  are  much 
alike  in  character.  This  species  is  much  smaller  than  0.  grandis.  The 
heads  are  of  about  the  same  size  but  not  alike  in  shape.  The  relations 
of  cirrus,  cirrus-pouch  and  vagina  are  different.    There  is  a  wide  differ- 


237]  PROTEOCEPHALIDAE—LA  RUE  237 

ence  in  the  number  of  testes  and  in  the  number  of  uterine  outpocketings. 
The  hosts  and  geographical  distribution  are  also  widely  different. 

It  varies  from  0.  marenzelleri  in  having  a  much  smaller  head, 
smaller  suckers  and  in  the  lesser  prominence  of  the  same,  in  its  smaller 
proglottids  and  in  its  smaller  number  of  testes.  The  relations  of  the 
cirrus  and  cirrus-sheath  are  quite  different.  The  fact  that  the  vagina 
lies  usually  posterior  to  the  cirrus-pouch  in  0.  marenzelleri  constitutes 
a  marked  difference.  The  shape  of  the  ovaries  is  somewhat  different. 
In  0.  marenzelleri  the  marked  bending  of  the  vagina  in  its  course  to 
the  middle  of  the  proglottid  is  very  unlike  the  condition  in  this  species, 
0.  calmettei  is  larger  than  0.  trimeresuri.  It  has  a  larger  head,  larger 
suckers  and  more  than  twice  as  many  testes.  The  relations  of  the 
unprotruded  cirrus  and  the  cirrus-pouch  are  much  alike  in  the  two 
species.  0.  calmettei  is  so  much  larger  than  0.  perspicua,  0.  nattereri 
or  0.  pigmentata  that  any  possibility  of  confusion  with  them  is  pre- 
cluded. It  is  likewise  much  larger  than  0.  filaroides  and  0.  lonnhergiii 
which  occur  in  amphibians.  0.  calmettei  most  nearly  resembles  0. 
grandis,  0.  ma,renzelleri  and  0.  trimeresuri  which  likewise  occur  in  the 
Crotalinae. 

OPHIOTAENIA  PUNIC  A  (Cholodkovski) 
[Figs.  153,  187,  188] 
Taenia  punica  Cholodkovski      1908:418-20 


1908 
1910 
1911 


Proteocephalus  punicus     Hall  1910:148 

Ophiotaenia  punica  La  Rue  1911 :481 


Specific  Diagnosis:  Characters  of  genus.  Length  as  much  as  10 
cm.  Breadth  up  to  2.75  mm.  Head  large,  1.5  mm.  broad,  unarmed, 
somewhat  tetragonal  in  shape.  Suckers  four,  large,  round,  0.7  mm.  in 
maximum  diameter.  Sucker  musculature  heavier  on  inner  half.  Necft 
short.  Segmentation  not  evident.  Proglottids  attached  by  full  width. 
First  proglottids  broader  than  long,  older  ones  quadrate  and  finally 
longer  than  broad.  The  genital  opening  alternates  irregularly,  is  situate 
near  middle  of  proglottid  margin.  Testes  very  numerous,  about  200, 
situated  in  two  well  defined  lateral  fields.  Cirrus-pouch  slender,  ex- 
tending 1/4  to  Ys  across  the  breadth  of  the  proglottid.  Lobes  of  ovary 
long,  slender,  rough  in  outline,  connected  by  a  long  slender  mid-piece. 
Vitellaria  lateral,  composed  of  small  follicles.  Uterus  median.  Eggs 
not  observed. 

Cholodkovski  (1908:418-420)  reported  this  species  from  a  dog  in 
Tunis  (Island  of  Dscherba).    Since  this  is  a  peculiar  host  in  which  to 


238  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [238 

find  a  species  normally  occurring  in  a  lower  vertebrate  his  description 
is  here  quoted  omitting  only  the  first  and  last  paragraphs: 

"Nicht  ohne  tlberraschung  habe  ich  also  in  einer  mir  von  Dr.  M.  Weinberg 
zugesandten  Tanie  aus  dem  Darme  eines  Hundes  eine  neue  Species  erkannt,  die 
ich  hier  in  aller  Kurze  beschreiben  will.  Das  Material  (in  Formol  aufbewahrt) 
stammte  aus  Turtis  (Insel  Dscherba)  und  bestand  aus  drei  unvollstandigen  Stro- 
biiae,  deren  grosste  eine  Lange  von  etwa  lo  cm  erreichte.  Der  Scolex  ist  sehr 
gross  (1.5  mm  breit),  unbewaflFnet,  mit  vier  starken  rundlichen  Saugnapfen  und 
einer  kleinen  Erhohung  auf  dem  Scheitel  versehen.  Die  Saugnapfe  sind  etwas 
asymmetrisch  gebaut,  indem  ihre  innere  Halfte  merklich  dicker  als  die  aussere  ist; 
der  grosste  (aussere)  Durchmesser  derselben  betragt  0.7  mm.  Gleich  hinter  einem 
sehr  kurzen  Halse  wird  die  Strobila  etwa  2  mm.,  dann  bis  2.75  mm.  breit,  nach 
hinten  aber  allmahlich  etwas  enger.  Eine  aussere  Gliederung  ist  auf  ganzen 
Stucken  unbemerkbar  und  lasst  sich  erst  auf  gefarbten  Balsam— oder  Glyzerin- 
praparaten  in  der  Gestalt  von  sehr  schwach  ausgesprochenen  Querrinnen  kon- 
statieren.  Die  ersten  Proglottiden  sind  breiter  als  lang,  (vide  Fig.  188  which  is 
reproduced  from  Cholodkovski's  figure  3)  dann  werden  sie  allmahlich  langer, 
qu^jdratisch  und  endlich  langer  als  breit;  in  keinem  der  mir  vorgelegenen  Exem- 
plare  waren  aber  ganz  reife;  d.  h.  fertige  Eier  enthaltende  Proglottiden  vorhanden. 
Die  Geschlechtsoffnungen  alternieren  unregelmassig.  Die  stark  in  der  Querricht- 
ung  verlangerten  Ovarien  liegen  dicht  an  der  hinteren  Grenze  der  Proglottis,  der 
Dotterstock  ist  sehr  klein,  der  Uterus  bildet  einen  geraden,  in  der  Mittellinie  der 
Proglottis  nach  vorn  verlaufenden  Stamm.  In  voUig  reifen  Proglottiden  treibt 
er  vielleicht  auch  laterale  Zweige,  in  meinen  Exemplaren  war  er  aber  immer 
einfach  (vide  Fig.  187  reproduced  from  Cholodkovski's  figure  4).  Die  zahlreichen 
Hoden  liegen  in  zwei  lateralen  Langsfeldern  nach  innen  von  den  grossen  Excre- 
tionsgefassen.  An  der  Stelle  letzteren  befindet  sich  in  totalen,  mit  Boraxkarmin 
tingierten  Praparaten  der  reifsten  mir  vorgelegnen  Proglottiden  eine  kornige 
Masse,  deren  Bedeutung  mir  unklar  geblieben  ist,  da  ich  aus  Mangel  an  Material 
keine  darauf  bezuglichen  Schnittserien  verfertigen  konnte. 

"Da  die  hier  beschriebene  Tanie  allem  Anschein  nach  eine  neue  Species 
darstellt,  so  schlage  ich  vor,  sie  nach  ihrem  Fundorte  Taenia  punica  zu  nennen." 

This  species  was  overlooked  by  the  writer  until  his  attention  was 
called  to  it  by  Hall's  article  (Hall  1910)  in  which  it  is  stated  that  the 
species  belongs  to  the  genus  Proteocephalus,  The  paragraphs  (Hall 
1910:146  and  148)  in  which  he  gives  his  reasons  for  this  belief  are  here 
quoted: 

"In  compiling  a  key  to  the  dog  tapeworms,  an  examination  of  Kholodkovski's 
(1908)  description  and  figures  of  Taenia  punica  from  the  dog  showed  that  the 
cestode  in  question  probably  belongs  in  the  genus  Proteocephalus  Weinland.  The 
head,  the  uterine  stem,  the  position  of  the  ovaries  at  right  angles  to  the  uterine 
stem,  and  the  position  of  the  testes  and  the  genital  canals  all  indicate  this.  The 
granular  strand  of  uncertain  nature  which  Kholodkovski  noted  in  the  position  of 


239]  PROTEOCEPHALIDAE—LA  RUE  239 

the  excretory  canals  can  hardly  be  anything  other  than  the  vitellaria,  in  the  loca- 
tion usual  for  the  species  of  the  genus  Proteocephalus.  Kholodkovski  states  that 
the  vitellarium  is  very  small,  but  it  seems  likely  that  he  has  mistaken  something 
else  for  the  vitellarium.  A  comparison  of  the  figures  with  mounted  specimens  of 
the  worms  of  the  genus  Proteocephalus  leaves  no  reasonable  doubt  on  this  point, 
and  it  is  the  opinion  of  the  writer  and  Dr.  B.  H.  Ransom,  with  whom  the  point 
was  discussed,  that  it  is  more  likely  that  the  dog  from  which  the  tapeworms  were 
obtained  had  just  eaten  the  true  host,  some  fish,  reptile,  or  batrachian,  than  that 
the  dog  was  the  true  host  by  virtue  of  a  normal,  even  though  unusual,  infection 
with  larval  form.  Fuhrmann  appears  to  have  overlooked  the  unusual  features  of 
this  worm  in  his  review  of  Kholodkovski   (1909),  and  states  that  the  anatomy  is 

that  of  species  of  Taenia." 

"Taenia  punica  Kholodkovski,  1908,  should  therefore  be  known  as  Proteoceph- 
alus punicus  (Kholodkovski,  1908)  Hall,  1910,  a  combination  proposed  here  for 
the  first  time " 

La  Rue  (1911:481)  included  this  species  in  the  genus  Ophiotaenia. 

After  making  a  careful  study  of  Cholodkovski 's  description  and 
drawings,  and  after  comparing  them  with  many  specimens  of  Proteo- 
cephalidae  the  writer  agrees  with  Hall  that  the  normal  host  of  this 
species  is  not  the  dog.  In  the  writer's  opinion  the  true  host  is  a  snake. 
This  cestode  having  its  testes  in  two  fields  does  not  resemble  any  of  the 
species  of  Proteocephalus  thus  far  described  from  fish.  The  Proteoceph- 
alids  that  infest  the  lizards  belong  to  the  genus  Acanthotaenia  and 
these  are  distinguished  from  other  Proteocephalids  by  the  presence  of 
spines  on  the  head  and  neck.  The  Proteocephalids  thus  far  described 
from  Amphibia  are  small  with  rather  small  heads.  Their  genital  organs, 
however,  resemble  those  of  Taenia  punica.  It  is  also  true  that  in  this 
respect  they  agree  with  the  Proteocephalids  of  snakes.  The  species 
from  snakes  vary  in  size  from  small  to  large.  Some  of  the  species  have 
large  heads  with  large  and  prominent  suckers.  Their  genital  organs 
much  resemble  those  of  Taenia  punica.  As  in  the  latter  species  the 
vitellaria  of  the  Proteocephalids  of  snakes  are  sometimes  composed  of 
small  follicles  which  refuse  to  take  the  stain  well.  In  such  cases  it  is 
easy  to  overlook  the  vitellaria.  As  an  instance  of  this  might  be  men- 
tioned the  Taenia  eunectes  A.  J.  Smith  in  which  the  coils  of  the  ducts 
in  the  interovarial  space  were  thought  to  be  the  vitellaria.  For  these 
reasons  and  also  on  account  of  the  size  of  the  head  and  strobila  the 
writer  suggests  that  the  true  host  of  Taenia  punica  is  a  snake,  and  that 
Taenia  punica  belongs  to  the  genus  Ophiotaenia  and  should  therefore 
be  known  under  the  name  Ophiotaenia  punica  (Cholodkovski,  1908) 
La  Rue. 


240  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [240 


1898 
1908 
1911 


OPHIOTAENIA  MARENZELLERI   (Barrois) 

[Figs.  37,  199] 

Ichthyotaenia  marenzelleri         Barrois  1898 :2-3 

Ichthyotaenia  marenzelleri         Schwarz         1908:26-27 
Ophiotaenia  marenzelleri  La  Rue  1911:481 


Specific  Diagnosis:  Characters  of  genus.  Observed  length  of  stro- 
bila  as  much  as  40  cm.  Maximum  breadth  1.5-2.0  mm.  Scolex  large, 
round,  muscular,  1.2-2.0  mm.  broad,  well  set  off  from  neck.  No  rostel- 
lum.  No  hooks.  No  functional  fifth  sucker.  Suckers  four,  0.60-0.70 
mm.  in  diameter.  Neck  not  observed.  Strobilation  distinct.  Proglot- 
tids  closely  attached.  Length  of  proglottids  as  much  as  5.5  mm.,  maxi- 
•mum  breadth  of  same  1.5-2.0  mm.  Excretory  system  composed  of  two 
pairs  of  lateral  vessels.  Genital  organs  as  in  genus.  Genital  aperture 
marginal,  irregularly  alternating,  situated  near  middle  of  lateral  mar- 
gin of  proglottid.  Testes  numbering  150-200-240,  measuring  0.06-0.07 
mm.  in  diameter,  situated  in  two  lateral  fields.  Cirrus-pouch  large^ 
extending  about  one-third  across  the  proglottid  breadth.  Cirrus,  when 
protruded,  swollen  at  base  and  filiform  at  tip,  1  mm.  long.  Vagina 
usually  posterior  to  cirrus-pouch.  Uterus  when  completely  developed 
possessing  20-25  lateral  outpocketings  on  either  side.  Eggs  provided 
with  two  membranes.    Diameter  not  known. 

Habitat:  In  intestine  of  Ancistrodon  piscivorous  Holbr.,  southern 
United  States. 

Barrois  (1898)  in  a  very  brief  description,  unaccompanied  by 
drawings,  proposed  this  species.  Schwarz  (1908:26-27)  redescribed 
this  species  using  Barrois 's  material.  His  description  was  accompanied 
by  three  drawings.  La  Rue  (1911:481)  included  this  species  in  the 
new  genus  Ophiotaenia. 

The  material  which  Barrois  and  Schwarz  had  for  study  was  col- 
lected by  Doctor  Calmette,  Dec.  22,  1897,  from  Ancistrodon  piscivorous 
Holbr.,  a  snake  indigenous  to  the  southern  United  States.  Unfortu- 
nately specimens  of  this  species  could  not  be  had  by  the  writer  for 
study  hence  the  data  used  in  the  following  description  are  derived  only 
from  the  articles  of  Barrois  (1898)  and  Schwarz  (1908).    A  study  of 


241]  PROTEOCEPHALIDAE—LA  RUE  241 

new  material  is  desirable  in  order  that  a  more  complete  knowledge  of 
this  form  may  be  had. 

This  is  one  of  the  largest  of  the  species  of  Ophiotaenia.  The  ob- 
served length  is  about  40  cm.  Schwarz  (1908:26)  thought  that  perhaps 
the  length  might  exceed  that  of  0.  calmettei  (35-80  cm.).  The  maxi- 
mum breadth  observed  was  1.2-2.0  mm.  The  scolex  is  large,  round  and 
muscular.  It  has  a  breadth  of  1.5-2.0  mm.  Barrois  described  an  un- 
armed rostellum.  Schwarz  found  no  rostellura,  nor  does  his  delineation 
of  the  head,  which  is  reproduced  (Fig.  37),  show  such  a  structure.  The 
scolex  is  clearly  set  off  from  the  neck.  The  four  suckers  are  muscular, 
prominent  and  nearly  circular.  They  measure  0.60-0.70  mm.  No  func- 
tional fifth  sucker  is  present.  The  neck  was  not  described  by  Barrois 
or  by  Schwarz.  The  strobilation  is  distinct,  altho  the  strobila  has  no 
indentations.  Proglottids  are  strongly  attached.  The  length  of  the 
largest  proglottids  is  about  5.5  mm.  and  the  maximum  breadth  about 
1.5-2.0  mm.  The  excretory  system  is  made  up  of  two  pairs  of  lateral 
vessels,  a  dorsal  and  a  ventral. 

The  genital  aperture  is  marginal  and  it  alternates  irregularly.  It 
is  situated  about  the  middle  of  the  proglottid.  The  testes  (Fig.  199) 
lie  in  two  well  marked  lateral  fields  between  the  vitellaria  and  the 
uterus.  Barrois  gives  230-240  as  their  number,  Schwarz  150-200.  They 
measure  0.060-0.070  mm.  Of  the  cirrus-sheath  and  vas  deferens 
Schwarz  says  "the  relation  of  the  cirrus-sheath  and  vas  deferens  is 
especially  characteristic  for  this  form.  In  ripe  segments  the  greater 
part  of  the  cirrus-sheath  is  completely  evaginated.  There  seems  to 
exist  a  special  muscular  apparatus  which  withdraws  the  cirrus-sheath, 
when  completely  sexually  mature,  from  the  proglottid,  so  that  it  pro- 
jects free  from  the  margin.  In  young  segments  it  lies  in  its  normal 
position  within  the  segment". 

Schwarz  has  evidently  misinterpreted  the  appearance  in  this  case. 
His  own  drawings  show  no  evagination  of  the  cirrus-pouch  but  a  com- 
plete evagination  of  the  cirrus  which  is  thick  at  the  base,  and  filiform 
at  the  tip.  From  this  heavy  base  the  ductus  ejaculatorius  runs  straight 
through  the  now  greatly  shrunken  cirrus-pouch.  There  are  some  coils 
of  the  ductus  ejaculatorius  within  the  basal  part  of  the  cirrus  itself. 
The  writer  has  observed  this  condition  in  0.  trimeresuri.  Schwarz 
states  that  the  cirrus  is  about  1  mm.  long  in  0.  marenzelleri.  In  his 
drawing  the  cirrus-pouch  extends  about  %  across  the  proglottid  breadth. 
The  vas  deferens  forms  a  mass  of  coils  before  entering  the  cirrus-pouch. 

The  vagina  opens  usually  posterior  to  the  cirrus-sheath.  Its  course 
to  the  middle  of  the  proglottid  forms  an  arc.  At  the  middle  of  the 
proglottid  it  bends  sharply  posteriad  and  then  its  course  is  direct  to 


242  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [242 

the  interovarial  space.  The  uterus  is  a  median  tube  from  which  arise 
on  either  side  many  lateral  pouches;  20-25  are  figured  in  Schwarz's 
drawing.  The  ovary  is  relatively  large  and  its  lobes  are  somewhat 
wing-like  in  shape.  The  eggs  are  provided  with  two  membranes.  No 
processes  arise  from  the  outer  one.  No  measurements  of  the  eggs  were 
given  by  Barrois  or  Schwarz. 

The  species  is  clearly  differentiated  from  the  other  large  species  of 
the  genus  by  the  size  and  shape  of  the  head  and  suckers.  The  relations 
of  the  cirrus  and  cirrus-pouch  vary  greatly  from  the  plan  common  to 
the  other  large  species  of  the  genus.  0.  trimeresuri  which  has  the  same 
relations  of  cirrus  and  cirrus-pouch  is  considerably  smaller.  0.  grandis 
resembles  it  in  having  the  same  host  species  and  in  being  of  large  size. 
0.  grandis  has,  however,  a  very  much  smaller  head  which  is  not  readily 
distinguished  from  the  strobila.  It  has  much  smaller  suckers  and 
greatly  different  relations  of  cirrus  and  cirrus-pouch.  Its  cirrus  is 
short  and  thick.  It  also  has  very  many  more  uterine  outpocketings. 
0.  marenzelleri  is  not  to  be  mistaken  for  0.  racemosa,  0.  imttereri,  0. 
pigmentata,  or  0.  perspicua  on  account  of  its  much  larger  size.  0.  fla- 
roides  and  0.  lonnhergii  being  from  Amphibia  and  also  being  much 
smaller  cannot  be  confused  with  0.  marenzelleri. 


OPHIOTAENIA  GRANDIS  La  Rue 
[Figs.  38,  97-101] 
1911 :     Ophioiaenia  grandis        La  Rue        1911 :481 

Specific  Diagnosis:  Characters  of  genus.  Body  very  long  (frag- 
ments 200  mm.)  by  2.75-4.25  mm.  broad  in  region  of  ripe  proglottids. 
Neck  5-8  mm.  long.  First  proglottids  much  broader  than  long;  pro- 
glottids with  developing  sexual  organs  quadrate  or  nearly  so;  ripe 
proglottids  quadrate  or  much  longer  than  broad.  Head  large,  1.0-1.2 
mm.  broad  at  base  of  suckers.  No  fifth  sucker  or  rostellum,  no  hooks. 
Suckers  nearly  circular,  deep  and  muscular,  measuring  about  0.34  by 
0.36  mm.  Genital  aperture  marginal,  irregularly  alternating,  situated 
near  middle  of  proglottid.  No  genital  papilla  but  sometimes  a  deep 
depression  about  genital  pore.  Testes  large,  numerous,  200-250  in  num- 
ber, arranged  in  two  broad  lateral  fields.  Cirrus-pouch  0,24-0.26  mm. 
broad,  0.64-0.75  mm.  long,  length  equal  to  V^-Vs  of  proglottid  width. 
Cirrus  short  and  heavy.  Few  or  no  coils  of  ductus  ejaculatorius  in 
cirrus-pouch.  Vagina  anterior  or  posterior  to  cirrus-pouch.  Sphincter 
vaginae  heavy.    Uterus  with  40-60  lateral  outpocketings  on  either  side. 


243]  PROTEOCEPHALIDAE—LA  RUE  243 

Ventral  uterine  openings  2-8  in  number.  Outer  egg  membrane  ellip- 
soidal, 0.026  by  0.037  mm.  or  sometimes  spherical  0.032  mm.  in  diame- 
ter; second  egg  membrane  0.021-0.023  mm.  Embryos  0.015-0.016  mm. 
Outer  membrane  smooth. 

Type:  Specimens  in  collections  of  United  States  Bureau  of  Ani- 
mal Industry,  No.  14854. 

Habitat:  In  intestine  of  Ancistrodon  piscivorous  Holbr.  (type 
host).  National  Zoological  Park.  The  locality  from  which  this  host 
came  is  not  known.    The  species  is  found  only  in  North  America. 

La  Rue  (1911:481)  described  this  species  in  a  preliminary  way. 
Six  or  eight  heads  together  with  numerous  pieces  in  a  bottle  bearing  the 
U.  S.  Bureau  of  Animal  Industry  number  14854  were  labelled  "Ichthy- 
otaenia:  host  Ancistrodon  piscivorous;  location,  intestine;  locality  Nat. 
Zoo.  Park,  collected  S.  S.  Shawhan.  Nov.  18,  1907."  These  were 
secured  for  the  writer's  study  by  Professor  "Ward. 

No  complete  worm  was  found.  The  largest  piece  was  200  mm. 
long  and  3.5  mm.  broad  at  the  anterior  end  and  4.25  mm.  broad  at  the 
posterior  end.  This  piece  was  made  up  of  proglottids  that  were  well 
advanced  in  development,  being  nearly  mature  at  the  anterior  end  and 
nearly  ripe  at  the  posterior  end.  The  younger  proglottids  were  not 
distinctly  separated  by  furrows  but  the  more  mature  ones  showed  very 
evident  segmentation.  Judging  from  the  size  of  the  segments  and  their 
state  of  development  it  seems  likely  that  the  whole  worm  would  attain 
a  length  of  300-400  mm.  or  perhaps  even  more.  Like  other  species  of 
Ophiotaenia  the  body  is  quite  flattened.  However,  the  body  is  thicker 
than  has  been  observed  by  the  writer  in  0.  calmettei,  0.  nattereri,  0. 
trimeresuri,  0.  perspicua,  and  Crepidohothrium  gerrardii.  The  head 
(Fig.  38)  is  large  tho  not  so  large  as  in  0.  marenzelleri  (Barrois)  or 
Crepidohothrium  gerrardii  (Baird).  It  measures  1.0-1.2  mm.  in  diame- 
ter at  the  base  of  the  suckers.  The  scolex,  measured  from  its  tip  to  the 
base  of  the  suckers  has  a  length  of  0.50-0.60  mm.  There  is  no  rostellum, 
no  hooks,  no  fifth  sucker,  and  from  toto  mounts  no  vestigial  fifth  sucker 
may  be  observed.    As  yet  no  sections  of  the  head  have  been  made. 

The  four  deep  suckers  are  borne  on  the  broadest  part  of  the  head 
and  these  are  directed  outward.  They  are  not  prominent,  are  strongly 
muscular,  nearly  circular  in  outline  and  they  lack  the  upward  pointing 
projection  that  is  always  found  in  the  suckers  of  C.  gerrardii.  The 
suckers  measure  0.34-0.35  mm.  in  transverse  axis  by  0.35-0.36  mm. 
in  longitudinal  axis.  The  cavity  of  the  sucker  is  deep.  Its  open- 
ing measures  0.15-0.17  mm.  in  longitudinal  diameter  by  0.20-0.22  mm. 
in  transverse  diameter.     Immediately  back  of  the  head  there  is  some- 


244  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [244 

times  a  large  inflation  (Fig.  38).  Since  this  is  not  always  present  it 
must  be  due  to  the  contraction  of  the  heavy  longitudinal  muscles  of  the 
upper  part  of  the  neck.  The  neck  itself  is  broad,  thin  and  long,  being 
0.85-1.0  mm.  broad  by  4-8  mm.  long. 

The  first  proglottids  are  short,  their  breadth  equalling  several  times 
the  length.  As  the  proglottids  become  older  the  shape  changes  from 
the  rectangular  to  nearly  quadrate  or  quadrate.  Proglottids  ranging  in 
age  from  nearly  mature  to  ripe  are  usually  longer  than  broad.  Those 
proglottids  which  show  the  beginnings  of  the  cirrus-pouch  and  vagina 
measure  0.5  mm.  long  by  1.0  mm.  broad  while  those  in  which  the  anlagen 
of  the  testes  are  appearing  measure  0.680  mm.  long  by  0.850  mm.  broad. 
Both  length  and  breadth  increase  greatly  as  the  segments  near  maturity. 
A  few  nearly  ripe  proglottids  are  broader  than  long,  measuring  3.25 
mm.  long  by  4.25  mm.  broad.  The  longest  proglottid  found  measured 
8  mm.  long  by  2.75  mm.  broad  while  many  proglottids  measure  3-5  mm. 
long  by  2-3  mm.  broad.  The  surface  of  the  body  is  somewhat  wrinkled, 
due  perhaps  to  having  lain  twisted  for  a  long  time  in  the  alcohol.  There 
seem  normally  to  be  no  furrows  either  longitudinal  or  transverse.  Pro- 
glottid limits  are  not  well  defined  in  young  proglottids  but  are  well 
defined  in  the  older  ones. 

The  genital  aperture  is  marginal,  irregularly  alternating.  Fre- 
quently the  aperture  occurs  on  the  same  side  in  several  successive  pro- 
glottids but  always  in  greater  numbers  on  the  left  side.  The  greatest 
number  of  genital  apertures  found  occurring  successively  on  the  left 
side  was  ten.  No  papilla  marks  the  genital  aperture.  Indeed  in  many 
proglottids  the  muscles  are  so  contracted  about  the  genital  aperture  as 
to  cause  a  deep  depression  (Fig.  100).  In  such  cases  it  is  noteworthy 
that  the  cirrus  and  vagina  open  to  the  exterior  separately.  In  many 
cases  it  is  doubtful  if  a  true  genital  sinus  exists.  If  present  it  is  ex- 
tremely shallow.  The  arrangement  of  the  sexual  organs  agrees  in 
general  with  that  of  other  Proteocephalids. 

The  cirrus-pouch  (Figs.  97,  98)  is  short  and  broad,  and  is  broadest 
at  the  distal  end.  It  is  either  straight  or  curved,  depending  upon  the 
state  of  contraction  of  the  proglottid  and  of  the  muscles  in  the  region 
of  the  genital  aperture.  When  the  region  of  the  genital  aperture  is 
depressed  the  cirrus-pouch  is  usually  somewhat  bent  (Fig.  100).  Its 
length  is  0.64-0.75  mm.  and  its  breadth  0.24-0.26  mm.  The  ratio  of  its 
length  to  the  breadth  of  the  proglottid  is  1 :3  or  1 :5.  The  cirrus  (Figs. 
97,  98)  when  protruded  is  thick,  short,  straight  and  heavily  muscled. 
It  is  continuous  with  the  ductus  ejaculatorius  which  forms  no,  or  at 
least  few,  coils  within  the  cirrus-pouch.  The  drawings  (Figs.  97,  98) 
show  only  a  slight  twist  or  turn  in  this  part  of  the  ductus.    This  descrip- 


245]  PROTEOCEPHALIDAE—LA  RUE  245 

tion  and  an  examination  of  the  figures  makes  it  evident  that  the  cirrus 
and  cirrus-pouch  of  the  present  species  are  very  different  from  the  same 
organs  in  0.  marenzelleri  with  which  this  species  is  closely  allied.  In 
that  species  there  are  many  coils  of  the  ductus  ejaculatorius  within  the 
cirrus-pouch,  and  the  cirrus  when  protruded  is  thick  at  the  base  but 
drawn  out  to  a  slender  tip.  Schwarz  (1908)  errs  in  saying  that  the 
cirrus-pouch  is  evaginated  in  0,  marenzelleri.  His  own  figures  show  the 
cirrus-pouch  in  situ.  In  the  present  species  the  vas  deferens  is  thrown 
into  numerous  coils  which  serve  as  a  vesicula  seminalis.  There  are  about 
200-250  testes  which  are  arranged  in  two  lateral  fields.  The  testes 
measure  0.04-0.05  mm.  broad  by  0.075-0.100-0.125  mm.  long.  The  vagina 
opens  to  the  exterior  either  anterior  or  posterior  to  the  cirrus-pouch, 
with  almost  equal  frequency.  Out  of  45  proglottids  examined  on  this 
point  the  vagina  in  21  cases  opened  anterior  and  in  24  cases  posterior 
to  the  cirrus-pouch.  A  heavy  sphincter  vaginae  0.095  mm.  long  by 
0.150  to  0.160  mm.  broad,  including  vagina,  surrounds  the  vagina  near 
its  opening  to  the  exterior.  The  thickness  of  the  sphincter  alone  is 
0.040-0.070  mm.  The  vitellaria  are  follicular  in  structure,  and  lateral 
in  position.  Each  follicle  measures  about  0.015  by  0.030-0.035  mm.  A 
diagram  (Fig.  99)  made  from  a  toto  preparation  shows  that  the  arrange- 
ment of  the  organs  of  the  interovarial  space  is  similar  to  that  figured 
for  other  Proteocephalids.  The  ovaries  are  thin  dorsoventrally.  The 
mid-piece  connecting  the  ovarian  lobes  is  slender  and  long.  The  uterus 
(Fig.  101)  in  maturing  proglottids  is  a  median  ventral  tube.  From  this 
are  developed  40-60  lateral  pouches  on  either  side.  These  are  formed 
by  the  method  already  described  by  La  Rue  (1909)  for  0.  filaroides. 
No  eggs  are  passed  into  the  uterus  until  the  outpocketings  have  attained 
a  considerable  size.  Two  or  more  preformed  ventral  uterine  pores  are 
developed  before  many  eggs  are  present  in  the  uterus.  The  largest 
number  of  the  uterine  pores  in  the  material  studied  was  eight.  The 
eggs  taken  from  the  uterus  of  alcoholic  specimens  are  usually  ellip- 
soidal, measuring  about  0.026  by  0.037  mm.  over  the  outer  membrane. 
Spherical  eggs  measure  about  0.032  mm.  The  second  membrane  is 
spherical,  0.021-0.023  mm.  in  diameter.  The  third  membrane  closely 
invests  the  embryo  which  is  0.015-0.016  mm.  in  diameter.  Mature  eggs 
have  smooth  contours.  There  are  no  booklets  or  holding  organs  on  the 
egg  membrane  as  Schwarz  (1908)  figures  for  the  eggs  of  0.  natter'eri 
(Parona). 

The  present  species  differs  from  0.  marenzelleri  (Barrois)  in  hav- 
ing a  smaller  head,  smaller  suckers,  larger  proglottids,  greater  breadth, 
more  numerous  testes  and  more  numerous  uterine  pouches.  The  char- 
acter of  the  evaginated  cirri  of  the  two  species  is  very  different.    It  also 


246  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [246 

differs  from  Crepidobothrium  gerrardii  (Baird)  in  the  size  of  the  head 
and  suckers,  and  in  the  character  of  the  latter.  0.  grandis  has  a  long 
unsegmented  neck  while  C.  gerrardii  has  almost  no  neck.  0.  grandis 
somewhat  resembles  0.  calmettei  in  the  size  of  the  head  and  suckers.  It 
differs  from  that  species  in  the  size  of  strobila  and  of  proglottids,  and 
in  length  of  neck.  0.  grandis  has  a  larger  number  of  testes  and  a  cirrus 
which  differs  greatly  from  that  of  0.  calmettei.  0.  grandis  is  so  much 
larger  than  0.  perspicua,  0.  nattereri  (Parona),  0.  irimeresuri  (Parona), 
and  0.  racemosa  (Rudolphi)  that  a  possibility  of  confusing  it  with  any 
of  those  species  is  precluded. 


CREPIDOBOTHRIUM  GERRARDII  (Baird) 

[Figs.  12,  13,  33,  34,  42,  111-115,  123,  124,  190,  196] 

1850:  Taenia  racemosa  Diesing  1850:511,  in  part 

1860:  Tetralothrium  gerrardii  Baird  1860:446-448 

1861:  Tetralothrium  gerrardii  Baird  1861:228-230 

1864:  Tetrahothrium  gerrardii  Diesing  1864:82 

1898:  Ichthyotaenia  gerrardii  Liihe  1898:652 

1899:  Crepidobothrium  gerrardii  Monticelli  1899:9-25 

1899:  Ichthyotaenia  gerrardii  Liihe  1899:525 

1905:  Taenia  racemosa  Shipley  1905:101 

1908:  Taenia  eunectes  Smith  1908:39-41 

1911 :  Crepidobothrium  gerrardii  La  Rue  1911 :479-480 

Specific  Diagnosis:  Characters  of  genus.  Length  of  strobila  as 
much  as  45.6  cm.  Maximum  breadth  0,85-2.00  and  even  6.0  mm.  Stro- 
bila flat,  thin  and  wrinkled.  Proglottids  closely  attached.  Segmentation 
indistinct.  Head  large,  tetragonal,  pyramidal,  maximum  breadth  of 
same  0.935-1.3  or  even  as  much  as  1.75  mm.,  thickness  a  little  less  than 
breadth,  length  about  0.68-0.90  mm.  Fifth  sucker  vestigial  but  with 
cavity  open  to  exterior.  Four  suckers  prominent,  muscular,  cordate, 
with  lower  margin  interrupted  and  re-entrant  into  sucker  cavity. 
Breadth  of  suckers  0.40-0.70  mm.  Neck  broad,  not  over  0.5-0.8  mm. 
long.  First  proglottids  short,  rapidly  increasing  in  length.  Mature 
proglottids  quadrate.  Ripe  proglottids  longer  than  broad.  Length  of 
same  1.7-2.3  mm.  by  0.65-1.09  mm.  broad. 

Genital  pore  marginal,  irregularly  alternating,  situated  near  mid- 
dle of  proglottid.  Vagina  anterior  or  posterior  to  cirrus-pouch.  Open- 
ing of  vagina  dorsal  to  cirrus-pouch.  Testes  200-400  in  number,  small, 
0.035-0.050-0.080  mm.  in  diameter,  situated  in  two  lateral  fields.   Cirrus- 


247] 


PROTEOCEPHALIDAE—LA  RUE 


247 


pouch  0.23-0.5  mm.  long  by  0.10-0.15  mm.  broad.  Length  of  same  4-5-6 
times  into  the  proglottid  breadth.  Several  coils  of  ductus  ejaculatorius 
in  cirrus-sheath.  Lumen  of  vagina  large  near  opening.  Sphincter 
vaginae  heavy.  Receptaculum  seminis  present.  Ovary  posterior,  bi- 
lobed,  lobes  thick,  short,  irregular.  Organs  of  interovarial  space  as  in 
Ophiotaenia.  Vitellaria  small,  loosely  follicular,  lateral.  Fully  devel- 
oped uterus  provided  with  20-30  lateral  pouches.  Eggs  possessing  twb 
(?)  or  three  membranes.  Outer  membrane  measuring  0.085-0.100  mm., 
second,  0.028-0.030  mm.,  embryo  0.016-0.018  mm.  in  diameter. 


Habitat : 
snakes. 


In  intestine  of  Boidae,   a  family  of  South  American 


Host 


(  ?)Ophiomorphus  miliaris 
Eunectes  murinus  (scytale) 

(?)Bothrops  jararacca 

i?)Ophis  merremii 
Boa  constrictor 
Eunectes  murinus  Wagl. 

Eunectes  murinus 


Locality 


South  America 
South  America 
South  America 
Brazil 
South  America 


Collector 


Natterer 
Edward  Gerrard 


A.  J.  Smith 


Authority 


Diesing,  1850:511 
Diesing,  1850 :5ri 
Diesing,  1850:511 
Diesing,  1850:511 
Baird,  1860:446-448 
Shipley,  1905:101 

(La  Rue) 
Smith,  1908:39-41 

(La  Rue) 


In  the  discussion  of  the  hosts  of  Ophiotaenia  racemosa  it  was  stated 
that  Ophiomorphus  miliaris  and  Ophis  merremii  were  to  be  considered 
as  probable  hosts  of  that  species.  It  is  doubtful  if  they  serve  as  the 
hosts  of  Crepidohothrium  gerrardii.  Bothrops  jararacca  is  one  of  the 
Crotalinae  and  would  not  be  likely  to  harbor  C.  gerrardii,  but  in  aH 
probability  it  would  be  the  host  of  a  species  resembling  0.  marenzelleri, 
0.  grandis,  0.  trimeresuri,  and  0.  calmettei.  These  form  a  group  quite 
distinct  from  the  other  Ophiotaenia.  It  is  very  likely  that  Diesing 
being  unable  to  make  a  careful  examination  of  the  cestodes  of  these 
snakes  grouped  them  all  under  the  name  Taenia  racemosa.  It  is  desira- 
ble to  re-examine  the  specimens  to  which  Diesing 's  description  and 
notes  refer,  if  they  still  exist.  The  hosts  of  C.  gerrardii  of  which  there 
is  no  doubt  are  Eunectes  murinus  "Wagl.  and  Boa  constrictor  L. 

This  species  was  first  reported  by  Diesing  (1850:511)  who  called  it 
Taenia  racemosa  Rud.  His  diagnosis  agrees  excellently  with  that  of 
Crepidohothrium  gerrardii.    He  includes  as  a  host  one  of  those  species, 


248  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [248 

Eunectes  murinus  (scytale),  from  which  this  species  has  been  reported 
several  times.    His  description  and  habitat  data  are  quoted  in  full: 

"Taenia  raccmosa  RUDOLPHI.  Caput  magnum  tetragonum,  acetabulis  an- 
gularibus  subterminalibus  v.  terminalibus,  subovatis  v.  cordatis.  Colluni  nullum. 
Articuli  supremi  brevissimi,  subsequentes  longiores  et  latiores,  angulis  rotundatis, 
ultimi  longi  parallelopipedi  angustiores.  Penes  fili formes  basi  incrassati,  margi- 
nales  vage  alterni.     Longit.  2"  il4' ;  latit.  med.  1-3"';  ultim.  i"'. 

"Taenia  racemosa  Rudolphi:  Synops.  App.  692. — Dujardin:  Hist.  nat.  des 
Helminth.  610. 

"Habitaculum.  Ophiomorphus  miliaris,  Martio. — Eunectes  Scytale,  Octobri, 
Februario  et  Junio. — Bothrops  Mararacca  (misspelling  for  jararacca),  Martio  et 
Aprili. — Ophis  Merremii,  Januario,  in  Brasilia  (Naiterer)  :  in  intestinis.  M.  C.  V." 

Baird  (1860:446-448)  described  this  species  which  was  found  by 
Mr.  Edward  Grerrard  in  the  intestine  of  a  Boa  constrictor  from  South 
America.  Since  the  name  Taenia  racemosa  Rud.  had  already  been  ap- 
plied to  a  different  species  of  cestode  the  name  suggested  by  Baird 
stands  by  reason  of  its  priority.  Baird 's  second  description  (1861:228- 
230)  is  a  copy  of  the  earlier  description  (1860:446-448).  Because  of 
the  inaccessibility  of  either  of  these  descriptions  for  many  investigators 
the  later  diagnosis  which  alone  was  accessible  is  quoted : 

" The    species   now    to   be   described,    however,    was    found   by   Mr. 

Edward  Gerrard  of  the  British  Museum  ...  in  the  intestine  of  a  Boa  constrictor 
from  South  America. 

"The  head  is  large,  tetragonal;  the  four  bothria  disposed  crosswise,  joined 
by  the  margins ;  each  of  them  large,  round,  and  having  on  one  side  a  strong  ridge. 
Body  depressed,  narrow,  articulated.  No  distinct  neck.  Anterior  extremity  of 
body  very  narrow,  and  the  articulations  there  are  extremely  small,  becoming 
larger  as  they  descend,  the  inferior  being  quadrangular  and  rather  large.  The 
margins  of  the  articulations  somewhat  annulated,  but  having  no  appearance  of 
genital  apertures.  The  head  is  about  three-fourths  of  a  line  broad ;  but  I  could 
not  discover  any  mouth.  Apparently  only  fragments  of  these  worms  were  ob- 
tained; but  some  of  these  detached  pieces  were  about  18  inches  in  length. 

"Hab.    Intestines  of  Boa  constrictor." 

Diesing  (1864:82)  placed  this  species  in  the  genus  Tetrahothrium 
but  with  some  reservation.  His  diagnosis  is  quoted  in  full  in  order  that 
a  comparison  of  it  can  be  made  with  his  diagnosis  of  Taenia  racemosa, 
Diesing  (1850:511) : 

"Tetrahothrium  Gerrardii  Baird.  Caput  magnum  tetragonum,  bothriis  quatuor 
cruciatim   oppositis   subcircularibus   magnis,   marginalibus   contiguis,   singulo   costa 


249]  PROTEOCEPHALIDAE—LA  RUE  249 

1.)     Os.  .  .  .    Collum  nullum.     Corpus  depressum  augustutn,  articulis   su- 

premis  angustissimis,  posterioribus  quadrangularibus.     Aperturae  genitalium  .... 
Longit.  fragment!  i8",  latit.  capit.  Ya". 

"Habitaculum.  Boa  Constrictor:  ex  America  meridionali,  in  intestinis  (Ger- 
rard)." 

A  comparison  of  the  diagnosis  just  quoted  and  the  earlier  one  by 
Diesing  (1850:511)  leaves  no  doubt  that  the  two  diagnoses  concerned 
the  same  species,  namely,  Crepidobothrium  gerrardii.  Liihe  (1898:652) 
included  this  species  in  a  list  of  Ichthyotaenia  from  snakes.  He  gave 
no  reason  for  considering  it  a  member  of  this  genus.  Fuhrmann  (1899 : 
864)  in  a  footnote  made  this  statement  concerning  this  species:  "Liihe, 
M.,  Oochoristica  nov.  gen.  Taeniadarum.  (Zoolog.  Anz.  1898.  No.  576) 
giebt  an,  dass  T.  Gerrardii  (Baird)  in  das  Genus  Ichthyotaenia  gehore; 
dies  ist  aber  wie  mir  Prof.  Monticelli  mitteilte  und  wie  ich  mich  selbst 
an  den  von  ihm  nach  den  originalen  angefertigen  Zeichnungen  uber- 
zeugen  konnte,  keineswegs  der  Fall.  Es  gehort  diese  Form  in  ein  beson- 
deres  Genus,  das  Prof.  Monticelli  demnachst  unter  dem  Namen  Crepi- 
bothrium  publizieren  wird."  During  the  same  year  Monticelli  (1899: 
9-25)  made  a  report,  accompanied  by  drawings,  upon  the  type  material 
of  this  species  which  was  in  the  British  Museum.  He  also  studied  some 
material  which  had  been  sent  to  him  by  Prof.  Fr.  Bell.  As  a  result  of 
this  study  he  erected  the  genus  Crepidobothrium  for  Baird 's  species 
alone.  Liihe  (1899:525)  in  a  footnote  discussed  the  position  of  this 
species  which  he  still  maintained  belonged  to  the  genus  Ichthyotaenia. 
His  ground  for  this  belief  was  that  the  head  and  suckers  which  are 
different  from  those  of  other  Ichthyotaenia  constitute  only  a  specific 
difference.  He  stated  that  the  name  Crepibothrium  (Fuhrmann  1899) 
was  a  nomen  nudum.  He  had  not  seen  Monticelli 's  article  and  so  was 
in  no  position  to  judge  Monticelli 's  work. 

Shipley  (1905:101)  made  the  following  report  of  this  species: 
''Taenia  racemosa  Rud.  Diesing,  Syst.  Helm.  I.  p.  511.  Many  speci- 
mens from  Eunectes  murinus  Wagl.,  it  has  also  been  described  from  the 
intestine  of  Bothrops  jararacca  "Wagl."  Thanks  to  Professor  H.  B. 
Ward  who  secured  them  for  study,  the  writer  has  been  able  to  examine 
some  of  Shipley's  specimens  from  the  host  Eunectes  murinus.  They 
were  probably  from  the  same  lot  which  Shipley  recorded  in  the  above 
quotation.  They  proved  to  belong  to  the  species  Crepidobothrium  ger- 
rardii (Baird).  Smith  (1908)  found  some  cestodes  in  an  anaconda, 
Eunectes  murinus.    He  described  them  under  the  name  Taenia  eunectes. 

"1)  "Each  of  them  large,  round  and  having  on  one  side  a  strong  ridge" 
Baird  1.  c."     (Baird  :i86o  or  i86i). 


250  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [250 

Upon  request  Doctor  Smith  sent  some  of  this  material  to  Professor 
Ward  who  has  very  kindly  permitted  the  writer  to  study  it.  Study  of 
this  material  has  shown  that  it  belongs  to  the  species  Crepidohothrium 
gerrardii.  La  Rue  (1911:479-480)  gave  the  synonymy  of  this  species 
briefly  and  stated  some  of  his  findings. 

A  comparison  of  the  diagnoses  of  Diesing  (1850:511  and  1864:82) 
with  the  description  by  Baird  (1861)  and  with  the  description  by 
Monticelli  (1899:9-25)  leaves  no  ground  for  doubt  that  these  investi- 
gators were  reporting  the  same  species.  In  certain  respects  the  descrip- 
tion by  Smith  (1908:39-41)  differs  radically  from  that  of  Monticelli. 
However  an  examination  of  Smith's  material  has  shown  that  Smith 
made  numerous  misinterpretations  which  are  the  cause  of  the  apparent 
discrepancies  between  his  work  and  Monticelli 's.  In  his  study  of  this 
form  the  writer  has  found  Monticelli  correct  in  the  main.  It  has  al- 
ready been  stated  that  Shipley's  material  belonged  to  C.  gerrardii. 

The  question  concerning  the  correct  name  for  this  species  must  be 
settled.  From  a  structural  point  of  view  this  species  cannot  be  retained 
in  the  genus  Taenia  nor  in  the  genus  Tetrabothrium.  Anatomically 
this  species  is  one  of  the  Proteocephalidae.  This  has  been  recognized 
by  Liihe  (1898),  Fuhrmann  (1899),  and  Monticelli  (1899).  The  name 
which  Fuhrmann  (1899:864)  gave  for  it  can  be  dismissed  as  a  nomen 
nudum.  Liihe  (1898  and  1899)  considered  that  the  species  belonged  to 
Ichthyotaenia.  It  has  already  been  shown  in  the  discussion  of  the  genus 
Proteocephalus  that  the  name  Ichthyotaenia  Lonnberg  (1894)  is  a  syn- 
onym of  the  name  Proteocephalus  Weinland  (1858)  and  hence  cannot 
be  retained.  The  latter  name  also  antedates  the  name  Crepidobothriura 
Monticelli  (1899)  and  on  that  account  should  be  used  in  place  of  Crepi- 
dobothrium  if  structurally  Baird 's  species  shows  sufficient  agreement 
with  Taenia  filicollis  (amhigua)  Rud.  the  type  of  Proteocephalus.  An 
examination  of  Monticelli 's  (1899)  description  and  figures  and  of  the 
writer's  description  with  its  accompanying  figures,  shows  conclusively 
that  this  species  does  not  agree  anatomically  with  the  type  of  Proteo- 
cephalus and  that  this  species  really  belongs  to  a  different  genus.  The 
only  available  name  is  Crepidohothrium  Monticelli  (1899)  which  is  not 
invalidated  by  the  name  Crepibothrium  as  used  by  Fuhrmann  and 
which  was  unaccompanied  by  a  diagnosis,  since  the  word  Crepibothrium 
is  not  a  homonym  of  Crepidohothrium.  Liihe 's  objection  therefore  is 
not  valid. 

One  bottle  of  9  specimens  with  heads,  labelled  ^^Ichthyotaenia 
Gerrardii  (Baird)  ?  aus  Eunectes  murinus  intestinum  Berlin  Aquarium" 
was  received  from  Dr.  Anton  Collin,  Berlin,  in  answer  to  a  request  from 
Professor  Ward.    This  is  now  No.  10.179  in  Professor  Ward 's  collection. 


251] 


PROTEOCEPHALIDAE—LA  RUE 


251 


One  bottle  of  5  specimens  with  heads  and  some  pieces,  labelled  ' '  Taema 
racemosa  Rud  from  Eunectes  murinus."  was  received  from  A.  E.  Ship- 
ley, Cambridge.  The  material  is  now  No.  08.472  in  Professor  Ward's 
collection.  One  bottle  of  3  specimens  with  heads,  labelled  ^'Taenia 
Eunectes  from  Anaconda,  sm.  Intestine,  Phila.  Zoo.  G.  939.  U.  P.  Path- 
Hist.  1694,"  was  received  from  A.  J.  Smith,  Philadelphia.  This  is  now 
No.  10.190  in  Professor  Ward's  collection.  Seven  slides,  one  of  them  con- 
taining a  head,  labelled  "No.  1858,  From  Boa  Constrictor  Box  53-20, 
1893,"  were  received  for  examination  from  the  Bureau  of  Animal  In- 
dustry, Washington. 

A  careful  study  of  the  specimens  thus  brought  together  showed  that 
they  belonged  to  the  same  species.  Since  in  each  lot  one  or  more  heads 
were  present  a  careful  study  of  this  organ  could  be  made.  For  this 
purpose  balsam  mounts  were  made  from  some  heads  while  others  were 
cleared  and  examined  in  glycerine.  On  account  of  the  small  amount  of 
material  the  section  method  was  not  used  except  in  a  single  instance 
The  results  of  this  study  are  best  shown  in  the  accompanying  table. 
The  heads  used  for  measurement  were  chosen  quite  at  random,  for  an 
examination  of  the  material  showed  that  a  tabulation  of  measurements 
of  all  heads  was  unnecessary.    The  dimensions  are  in  millimeters. 


Lot  number 

Source 
Preparation 

1858 
B.A.I, 
balsam 

08.472  from 
Shipley 
balsam 

unflattened 

08.472  from 

Shipley 

glycerine 

10.179  from 

Anton  Collin 

glycerine 

10.179  from 

Anton  Collin 

glycerine 

10.190  from 

A.  J.  Smith 

glycerine 

T.  eunectes 

Data  from 

Smith  (1908) 

Head 
Suckers 

breadth 

length 

breadth    of 

two 

breadth 

length 

1.360" 

o.68o± 

0.646 

0.646 

1. 000 

o.50o± 

0.935' 

o.68o± 

0.400 

0.459 

0.76s 

0.680 

1.090 

o.68o± 

0.646 

1. 190 

o.68o± 

0.544 

O.S44 

0.402 

0.850 

1.090 

o.68o± 

0.510 

1. 190 
0.850 
0.510 

0.544 
1. 190 
0.500 

1.300 

0.900* 

0.700 

Neck 

0.850 
0.500 

0.390 
0.500 

0.300 
1.200' 

a)  Head   showed  some  eflfect  of  flattening  in  mounting.     This  may  have  some 

effect  on  measurements  of  suckers  also. 

b)  Head  was  evidently  very  strongly  contracted  as  were  also  the  suckers. 

c)  Measurements  were  taken   from  Smith's  drawing  which  was  accompanied  by 

a  scale  of  magnification. 


A  study  of  the  above  table  shows  that  in  the  main  the  measure- 
ments agree  very  well.  Where  apparent  discrepancies  occur  they  are 
easily  accounted  for  by  states  of  contraction  and  compression.  In  every 
case  the  shape  of  the  suckers  is  the  same,  due  allowances  being  made 


252  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [252 

for  contraction  and  relaxation.  Every  head  of  the  four  lots  was  exam- 
ined in  this  respect  and  in  each  sucker  the  lower  margin  was  turned  in 
thus  ** forming  an  angular  intrusion  into  the  cavity  of  the  sucker"  as 
Smith  (1908)  described  it.  In  the  more  contracted  or  in  greatly  ex- 
panded suckers  this  condition  was  somewhat  difficult  to  make  out  but 
when  properly  cleared  as  in  glycerine  or  balsam  it  was  apparent.  A 
study  of  toto  preparations  of  the  proglottids  from  the  various  lots  fails 
to  disclose  any  good  characters  for  separating  them.  The  writer 
therefore  bases  the  following  description  upon  this  material  so  collected 
together,  referring  at  times  to  special  lots.  He  also  uses  the  descriptions 
by  Smith  (1908)  and  by  Monticelli  (1899)  as  sources  of  comparative 
data.  It  was  desired  to  study  the  slides  upon  which  Monticelli  based 
his  description  but  unfortunately  Professor  Ward  was  unable  to  secure 
them. 

This  is  one  of  the  largest  of  the  Proteocephalid  species  inhabiting 
snakes.  0.  calmettei  (Barrois),  0.  marenzelleri  (Parona),  0.  grandis  are 
of  about  the  same  size.  The  latter  even  exceeds  C.  gerrardii  in  some 
measurements.  Next  smaller  than  the  members  of  this  group  comes 
0.  racemosa  (Rud.).  A  comparison  of  the  table  at  the  end  of  this 
report  shows  the  size  relationships  of  these  species.  The  strobila  varies 
considerably  in  length.  According  to  Baird  (1860)  it  measures  45.6 
cm.  (18  inches).  Smith  (1908)  reports  specimens  95-100  mm.  long. 
Diesing  (1850:511)  gives  its  length  as  2  inches  to  11/^  feet.  Monticelli 
gives  no  data  on  this  point.  The  reported  breadth  varies  from  0.85 
mm.  in  some  short  specimens  examined  by  the  writer  to  1.8  mm.  in  the 
B.  A.  I.  slide  1858,  to  2.0  mm.  according  to  Smith,  and  2.0-6.0  mm. 
(1-3  lines)  in  the  middle  to  2  mm.  (1  line)  at  the  posterior  end,  accord- 
ing to  Diesing  (1850).  The  strobila  is  flat  and  relatively  thin.  The 
surface  is  more  or  less  thrown  into  folds.  The  proglottids  are  attached 
by  their  full  width.  The  segmentation  is  indistinct  in  the  anterior 
region  and  more  distinct  in  the  region  of  mature  and  ripe  proglottids. 
In  certain  strobilas  the  segmentation  is  more  distinct  than  in  others. 
It  is  never  as  clearly  marked  as  in  some  species  of  Proteocephalids. 

The  head  is  large,  readily  distinguishable  to  the  naked  eye.  Baird 
describes  it  in  these  words:  "The  head  is  large,  tetragonal;  the  four 
bothria  disposed  crosswise,  joined  by  the  margins."  Smith  (1908)  says 
of  it,  "The  head,  viewed  from  the  front  presents  a  crucial  appearance 
from  the  prominence  of  the  suckers,  measuring  transversely  across  the 
two  opposed  suckers  1.5  mm.  and  laterally  across  two  adjacent  suckers 

1.3  mm The  suckers,  thus  prominent,  form  the  rounded  arms 

of  the  crucial  frontal  picture,  each  sucker  being  globose  in  shape  and 
having  a  lateral  diameter  of  about  0.7  mm."     Smith's  figures  of  the 


253]  PROTEOCEPHALIDAE—LA  RUE  253 

hea4  are  reproduced  (Figs.  33,  34).  Monticelli's  (1899)  description  of 
the  head  reads:  "Come  si  vede,  il  capo  (Monticelli's  figures  of  the 
head  are  reproduced  in  figures  123,  124)  e  abbastanza  grande  e  ben 
distinto  dal  coUo  che  anteriormente  si  slarga  di  poco  come  pei  sostenerlo : 
esso  ha  I'aspetto  grossolano  di  un  porno  da  bastone  e,  come  si  rileva 
dalla  figura,  e  rigonfio  e  massiccio,  tetragonale,  a  forma  di  piramide 
tronca.  Anteriormente  ristretto,  cupuliforme,  terminato  nel  mezzo  da- 
un  indistinto  cocuzzoletto  apicale  conoide ;  posteriormente  slargato  e  v 
troncato  sporge  oltre  il  coUo  per  i  suoi  spessi  margini  rigonfi  e  presenta, 
nel  mezzo  di  ciascuno  dei  quattro  lati,  una  insenatura  molto  accentuata. ' ' 
The  head  is  always  tetragonal  and  somewhat  pyramidal.  This 
shape  may  be  varied  to  some  extent  by  the  states  of  contraction.  A 
head  which  is  relaxed  and  whose  suckers  are  expanded  may  have  a 
lobate  appearance  (Fig.  13).  Here  deep  grooves  may  be  seen  between 
the  suckers.  In  greatly  contracted  heads  (Fig.  12)  the  suckers  are 
drawn  closely  together,  the  furrows  between  the  suckers  are  almost 
obliterated  and  the  suckers  themselves  are  reduced  in  size.  In  such  heads 
the  suckers  are  seen  to  be  directed  forward  while  in  the  relaxed  head 
the  suckers  are  directed  outward.  These  figures  should  be  compared- 
with  the  drawings  of  heads  by  Smith  (1908)  and  Monticelli  (1899), 
which  have  been  reproduced  here  (Figs.  33,  34  and  Figs.  123,  124). 
These  figures  illustrate  the  various  states  of  contraction  which  one  meets 
in  a  study  of  considerable  material.  The  dimensions  of  the  head  are: 
breadth  0.935-1.30  mm.  in  the  writer's  material,  1.75  mm.  (Baird), 
thickness  nearly  equals  breadth,  length  about  0.68-0.90  mm.  The  apex 
of  the  head  is  slightly  elevated,  rarely  flat.  There  is  no  rostellum  and 
no  spines. 

From  toto  preparations  no  trace  of  a  fifth  sucker  could  be  found 
but  in  a  series  of  transverse  sections  of  the  head  a  small  sunken  fifth 
sucker  was  found  which  had  degenerated  until  it  had  lost  its  radial 
musculature  (Fig.  42).  The  sucker  still  possessed  its  opening  to  the 
exterior  which  measured  0.037  by  0.026  mm.  and  this  opening  led  into 
a  cavity  but  a  little  more  than  0.03  mm.  deep.  The  sucker  measured 
about  0.048  by  0.067  mm.  on  its  transverse  axes.  From  the  tip  it  ex- 
tended about  0.06  mm.  into  the  head.  This  tissue  was  in  bad  condition. 
It  took  the  stain  poorly.  Its  nuclei  were  not  visible  either  in  the  sucker 
or  in  the  other  parts  of  the  head.  Cell  outlines  were  indistinct  or  lack- 
ing. The  musculature  of  the  sucker  was  irregular  or  perhaps  no  longer 
visible.  There  were  however  certain  fibers  which  seemed  to  be  muscles. 
A  basement  membrane  was  clearly  visible  and  surrounding  this  on  the 
outside  were  circular  muscle  fibers  which  could  be  readily  distinguished 
by  focussing  on  successive  layers  of  tissue.    The  sucker  cavity  was  filled 


254  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [254 

with  a  granular  mass  of  unknown  origin.  The  writer  noted  a  similar 
granular  material  in  the  sucker  cavities  of  degenerating  suckers  of 
Ophiotaenia  filaroides.  This  sucker  stands  about  midway  between  the 
vestigial  sucker  of  0.  filaroides,  O.  lonnhergii  and  P.  amhloplitis  on  the 
one  hand  and  the  functional  fifth  sucker  of  P.  fallax,  P.  pinguis  and 
P.  percae  on  the  other.  Approximately  the  same  stage  of  degeneration 
is  shown  in  the  drawing  (Fig.  44)  of  a  degenerating  fifth  sucker  of  the 
plerocercus  of  O.  filaroides  as  is  shown  by  the  fifth  sucker  in  C.  gerrardii. 
Of  the  suckers  Monticelli  (1899)  writes: 

"Non  vi  sono  botridii,  ma  quattro  ventose,  grandi  ovoidali,  disposte  in  croce, 
secondo  il  loro  asse  tnaggiore,  ai  quattro  angoli  del  capo  ed  obliguamente  e  con- 
vergenti  verso  il  cocuzzoletto  apicale.  Esse  sono  approfondate  nello  spessore  del 
capo  e  ne  occupano  quasi  tuta  la  lunghezza;  fra  le  ventose  il  capo  si  in  fossa  in 
piccoli  solchi  longitudinal  che  si  terminano  nelle  insenature  marginali  innanzi 
ricordato.  Le  ventose  hanno  forma  di  ferro  di  cavalloe,  diro  meglio,  ricordano  la 
figura  di  un  piede  di  cavallo  ferrato  visto  di  sotto.  Esse  hanno  un  aspetto  carater- 
istico  proprio,  che  puo  facilmente  ricavarsi  dalla  fig.  2  e,  meglio  ancora,  dalla  fig. 
3,  che  mostra  il  capo  visto  di  sopra  ed  alquanto  schiacciato.  Che  il  margine 
posteriore  delle  ventose  non  e  integro,  ma  scavato  da  una  insenatura  formata  dal 
ripiegarsi  verso  il  mezzo  del  cavo  di  esse  delle  pareti  posteriori  della  ventosa — 
che  non  formano  un  tutto  continuo,  ma  sono  posteriormente  interrotte — che  rav- 
vicinate  fra  loro,  constituiscono  una  doppia  cresta,  breve,  la  quale  si  perde,  divari- 
cando  in  due  rametti  a  V,  nel  fondo  della  ventosa.  Ma,  meglio  che  dalla 
descrizione.  puo  intendersi  la  peculiare  struttura  di  queste  ventose,  gia  adombrata 
dal  Baird,  come  si  rileva  dalle  parole  innanzi  citate,  dall'  esame  delle  figure  2  e  3, 
che  danno  una  fedele  immagine  della  forma  del  capo  del  cestode  in  esame,  come 
I'ho  ricavata  dallo  studio  del  tipo  originale  del  Baird." 

Monticelli 's  figures  above  mentioned  are  reproduced  (Figs.  123- 
124).  In  these  figures  the  point  of  the  in  turned  margin  is  split  and 
each  arm  of  the  "V"  is  turned  out.  Material  examined  by  the  writer 
shows  no  such  spreading  of  the  point  of  the  "V".  A  series  of  tran- 
sections show  that  the  point  of  the  '*V"  is  closed.  When  the  sucker  is 
much  relaxed  the  point  is  difficult  to  see,  but  it  may  be  found  if  the 
head  be  cleared  and  examined  in  glycerine.  Likewise  in  very  strongly 
contracted  suckers  the  intumed  point  may  escape  detection  except 
when  cleared  and  examined  under  favorable  conditions.  Prom  an  ex- 
amination of  three  heads  from  Smith's  material  the  writer  is  led  to 
believe  that  the  narrow  slit-like  opening  in  the  suckers  as  shown  in  his 
figures  (reproduced  Figs.  33,  34)  is  due  to  foreshortening.  Usually  the 
opening  is  much  more  nearly  round  than  here  shown.  The  suckers  are 
very  prominent,  most  prominent  in  a  relaxed  head  (Fig.  13).  They 
measure  0.40-0.70  mm.  in  breadth.    The  average  is  about  0.55  mm. 


255]  PROTEOCEPHALIDAE—LA  RUE  255 

A  poorly  defined  neck  region  separates  the  head  from  the  body. 
Baird  (1860)  says:  "No  distinct  neck.  Anterior  extremity  of  the 
body  very  narrow."  Monticelli  (1899)  says:  "II  collo  e  relativamente 
lunghotto  e  le  prime  proglottidi  sono  come  delle  rughe  e  molto  indis- 
tinte."  A.  J.  Smith  (1908)  gives  a  width  of  0.300  mm.  for  the  neck 
and  his  drawing  shows  a  length  of  1.20  mm.  Measurements  of  Smith's 
material  however  gives  a  width  up  to  1.190  mm.  Measurements  of  six 
strobilas  give  0.5-0.85  mm.  for  the  length  of  the  neck,  and  in  these  cases 
it  is  probable  that  the  measurements  given  are  too  great.  The  region  of 
proglottid  formation  begins  very  close  to  the  head  and  here  the  proglot- 
tids  are  very  short  and  their  boundaries  poorly  defined.  The  neck  is 
thin  and  flat.  The  first  proglottids  are  much  broader  than  long.  As 
they  become  older  they  increase  rapidly  in  length.  Mature  segments 
are  nearly  quadrate  while  ripe  proglottids  are  longer  than  broad.  The 
moniliform  proglottids,  mentioned  by  Smith  (1908),  are  not  character- 
istic of  the  species  but  are  due  to  contraction  states.  Among  the  three 
specimens  which  he  sent  to  Professor  "Ward  was  a  short  one  which  had 
these  peculiarly  contracted  proglottids.  The  longest  and  most  perfect 
specimens  had  no  segments  of  this  character  nor  were  such  proglottids 
observed  among  the  specimens  of  the  other  lots.  Ripe  proglottids  meas- 
ure 1.70-2.30  mm.  in  length  by  0.65-1.09  mm.  in  breadth. 

The  genital  anlagen  appear  very  early  in  the  chain.  In  fact  they 
are  to  be  seen  in  all  but  the  youngest  proglottids.  The  genital  aperture 
is  irregularly  alternating.  It  is  situated  near  the  middle  of  the  margin 
of  the  segment.  The  vagina  and  the  cirrus  open  into  the  common  geni- 
tal sinus,  the  vagina  being  either  anterior  or  posterior  to  the  cirrus- 
pouch.  A  genital  sinus  can  scarcely  be  said  to  exist.  Under  favorable 
circumstances  the  sinus  may  be  seen  to  be  0.020-0.030  mm.  deep.  This 
shallow  sinus  is  easily  overlooked.  The  opening  of  the  vagina  is  some- 
times dorsal  to  the  cirrus-pouch  but  the  writer  is  not  prepared  to  say 
that  it  always  occupies  that  position.  Monticelli  (1899)  thinks  that  the 
vagina  normally  lies  above  the  cirrus-pouch  and  that  when  it  is  found 
anterior  or  posterior  to  the  same  it  has  been  displaced  by  pressure. 
From  work  on  this  species  and  also  on  several  other  species  in  which 
the  vagina  may  lie  in  either  the  anterior  or  posterior  position  the  writer 
is  convinced  that  pressure  has  nothing  to  do  with  this  variation.  The 
preparations  studied  have  all  been  made  without  compression,  yet  the 
vaginae  alternate  irregularly  in  position.  Sections  in  a  frontal  plane 
through  developing  proglottids  of  0.  lonnbergii  show  the  vaginae  some- 
times anterior  and  sometimes  posterior  to  the  cirrus-pouch.  They  de- 
velop in  those  positions  which  they  hold  in  mature  and  ripe  proglottids. 
A  careful  examination  of  toto  preparations  of  C.  gerrardii  and  of  sev- 


256  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [256 

eral  species  of  Ophiotaenia  shows  that  if  the  vagina  is  in  an  anterior 
position  it  extends  farther  anteriad  before  bending  toward  the  genital 
pore  than  it  does  when  it  occupies  the  posterior  position.  If  these 
vaginae  are  displaced  by  pressure  then  they  ought  to  show  evidence  of 
such  displacement  by  their  form  or  structure.  This  they  do  not  do,  as 
an  examination  of  the  drawings  (Figs.  111-115)  will  show.  It  will  be 
noted  that  when  the  vagina  is  anterior  to  the  cirrus-pouch  it  usually 
crosses  the  coils  of  vas  deferens.  In  C.  gerrardii  the  vagina,  when 
anterior,  never  crosses  the  cirrus-pouch. 

The  testes  lie  in  two  broad  fields  which  tend  to  coalesce  in  the 
anterior  region  of  the  proglottid.  Testes  number  about  240  in  the 
specimens  from  the  Bureau  of  Animal  Industry.  Montieelli  (1899) 
found  about  200  or  more  in  each  proglottid.  Smith  (1908)  figures  but 
87  testes.  There  must  be  an  enormous  variation  in  the  number  of  testes 
or  else  Smith  overlooked  some  of  them,  for  in  a  preparation  of  an 
immature  proglottid  from  one  of  his  specimens  the  writer  counted 
nearly  400  testes.  In  this  proglottid,  delineated  (Fig.  113),  the  testes 
are  very  small.  In  maturity  many  of  them  might  be  crowded  down  to 
a  lower  plane  and  so  might  be  overlooked.  However,  in  his  prepara- 
tions the  writer  has  not  found  evidence  for  this  crowding.  There  is  no 
evidence  that  any  of  the  testicular  anlagen  fuse  together  or  degenerate. 
As  a  rule  the  testes  in  this  species  measure  0.030-0.050  mm.  in  diameter 
but  in  a  specimen  from  lot  10.179  the  testes  measured  as  much  as  0.080 
mm.  Montieelli  (1899)  says  that  they  are  about  twice  the  size  of  the 
vitelline  follicles.  The  vas  deferens  (Fig.  112)  in  ripe  proglottids  is 
thrown  into  numerous,  heavy,  spermatozoa-laden  coils  which  extend  in 
a  mass  nearly  to  the  middle  of  the  proglottid.  Within  the  cirrus-pouch 
the  ductus  ejaculatorius  makes  a  few  coils  before  passing  over  into  the 
cirrus.  The  latter  when  protruded  (Fig.  114),  is  short,  thick  and  very 
muscular.  Montieelli 's  description  agrees  with  this  statement.  The 
cirrus-pouch  is  short,  broad  and  heavily  muscled.  Its  length  is  about 
0.23-0.25  mm.  and  its  breadth  about  0.10-0.15  mm.  In  maturing  pro- 
glottids the  ratio  of  its  length  to  the  segment  breadth  is  1 :5  or  1 :6 
while  in  ripe  proglottids  it  is  about  1 :4. 

These  organs  (Figs.  112-115)  are  arranged  as  in  the  Ophio- 
taenia. The  lumen  of  the  vagina  (Figs.  114,  190)  near  its  opening  to 
the  exterior  is  large.  The  vagina  is  here  surrounded  by  a  strong  sphinc- 
ter vaginae.  There  are  no  coils  of  vagina  anterior  to  the  ovary.  A 
receptaculum  seminis  has  not  been  seen.  Montieelli  (1899)  states  that 
it  is  present.  The  ovary  is  bilobed  and  it  is  situated  in  the  posterior 
part  of  the  proglottid.  The  lobes  are  thick,  broad  and  short.  Their 
outlines  are  more  or  less  irregular.    An  oocapt,  ootype,  shell-gland,  ovi- 


257]  PROTEOCEPHALIDAE—LA  RUE  257 

duct,  unpaired  vitelline  duct,  lower  vagina,  and  a  uterine  passage  are 
present  and  these  organs  are  arranged  in  the  manner  common  to  the 
Proteocephalidae.  Monticelli  (1899)  gives  a  long  and  detailed  descrip- 
tion of  the  arrangement  of  these  organs  upon  which  no  comments  are 
necessary.  Smith  (1908)  mistook  these  organs  of  the  interovarial  space 
for  vitellaria. 

The  vitellaria  are  lateral  loosely  follicular  masses  which  extend 
throughout  the  length  of  the  proglottid.  The  follicles  are  small,  and 
in  some  preparations  they  took  the  stain  poorly.  It  is  probable  that 
Smith  had  the  same  difficulty  in  staining  the  vitellaria  and  for  that 
reason  he  entirely  overlooked  them.  Monticelli  states  that  the  vitellaria 
lie  outside  of  the  inner  longitudinal  muscle  layer.  The  writer  has 
examined  nothing  but  toto  preparations  which  perhaps  are  not  trust- 
worthy for  the  determination  of  this  point.  Nevertheless  it  seems  that 
Monticelli 's  statement  is  the  result  of  a  misinterpretation  of  the  facts. 
The  uterus  (Figs.  Ill,  113,  114,  115)  is  a  median  tube  which  in  ripe 
proglottids  (Fig.  112)  has  from  20  to  30  lateral  outpocketings  on  either 
side.  As  these  pouches  fill  with  eggs  they  become  rounded  and  the 
septa  become  indistinct.  The  uterus  is  not  connected  with  the  lobes  of 
the  ovary  as  Smith  (1908)  figures  it.  His  figure  is  incorrect.  The 
pouches  are  more  numerous  and  more  closely  applied  to  each  other  and 
less  slender  than  he  shows  them.  Uterine  pores  were  not  visible  in 
material  examined  by  the  writer.  As  to  the  method  of  discharging  the 
eggs  Monticelli  (1899)  writes:  "Nelle  ultime  proglottidi  I'utero,  pregno 
a  rimpinzato  d'uova,  sporge  alquanto,  facendo  ernia  contro  le  pareti, 
sulla  faccia  ventrale  delle  proglottidi.  Non  ho  osservato  orifizio  esterno 
permante  dell'utero,  come  quelle  che  Kramer  ha  visto  in  alcuni  Tetraco- 
tylus  (p.  e.  T.  filicoUis)  [Monticelli  here  cites  PL  28  fig.  42  in  Kramer's 
paper  of  1892]  e  neppure  ho  constat©  determinarsi  uno  sbocca  prowi- 
sorio  per  dar  uscita  alle  uova:  penso,  forse,  I'uscita  delle  uova  all 'esterno 
avvenga,  come  in  altri  casi,  per  deiscenza." 

According  to  Smith  (1908)  the  six-hooked  embryo  measures  0.016- 
0.018  mm.  in  diameter,  the  two  membranes  0.028-0.030  mm.  and  0.085- 
0.100  mm.  respectively.  The  writer  was  unable  to  measure  or  to  study 
any  eggs  of  this  species  hence  he  is  not  able  to  comment  on  Smith's  data 
nor  upon  Monticelli 's  description  of  the  eggs  which  is  here  quoted : 

"Le  uova  uterine,  relativemente  piccole,  hanno  un  guscio  spesso  e  molto 
evidente :  contengono  gia  I'oncosfera  involta  in  una  veste,  od  invoglio,  che  come 
una  teca  a  netto  contorno  e  non  molte  spessa,  per  quanto  ben  distinta,  circonda, 
come  in  altri  teniadi,  I'embrione.  Ciascun  uovo,  con  il  relativo  guscio,  e  racchiuso 
in  una  capsula  grande,  sferoidale  e  di  diametro  assai  pivi  del  doppio  del  guscio, 


258  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [258 

omologa  analoga  a  quella  che  si  osserva  in  altri  cestodi  (capsule  uterine,  od 
ovariche  Auct.).  La  quale,  come  altrove  ho  dimostrato  [Monticelli,  Fr.  Sav. — 
Sulla  cosidetta  subcuticola  dei  Cestodi,  in :  Rend  R.  Ace.  So.  Napoli,  Fasc.  7-12, 
Luglio-Decembre  1892  (sul  princpio  dove  parlo  dell'uovo  dei  Cestodi  e  ne  riassumo 
lo  sviluppo).]  nulla  ha  da  vedere  col  guscio  delle  uova  essendo  essa  una  forma- 
zione  molto  diverso,  secondaria  e  posteriore :  circa  I'origine  di  questa,  oltre  quanto 
ho  espresso  nel  citato  mio  lavoro,  veggasi  pure  il  Diamare  [Diamare,  V. — II  genere 
Dipylidium,  in:  Atti  R.  Accad.  Sc.  Nap.  (2)  Vol.  VI.  N.  7,  31  pp,  3  Tavole]. 
Capsula  queste  che  ritengo  possa  e  debba  venir  interpretata  come  una  sorte  di 
cocon  e  corrispondente  a  formazioni  del  genere  che  si  osservano  nelle  uova  di 
altri  animali  e  ne  circondona  il  guscio." 

Since  Smith's  specimens  of  Taenia  eunectes  Smith  came  from  a  host 
which  is  not  uncommon  in  American  zoological  gardens  and  since  his 
paper  will  probably  be  widely  referred  to  by  American  investigators 
more  specific  attention  is  called  to  certain  misinterpretations  in  the 
description  and  delineation  of  this  species  by  Smith  (1908) .  His  figure  4, 
PI.  Ill  which  has  not  been  reproduced  shows  very  clearly  that  he  con- 
sidered this  form  to  be  a  species  of  Taenia.  His  attempt  to  make  the 
organs  of  this  species  agree  with  those  of  Taenia  has  resulted  in  consid- 
erable confusion.  The  organ  which  he  has  labelled  the  vagina  is  the 
cirrus-pouch  and  vice-versa.  The  receptaculum  seminis  of  his  labelling 
is  the  mass  of  coils  of  the  vas  deferens.  The  lateral  vitellaria  he  has 
omitted  entirely.  He  labels  vitellaria  that  mass  of  coils  comprising  the 
organs  of  the  interovarial  space.  The  ovary  in  this  species  is  never 
connected  with  the  uterus  as  he  figures  it,  but  always  through  the  single 
oviduct,  ootype,  and  uterine  passage  as  has  been  figured  (Figs.  99,  104). 
The  writer  has  never  seen*  the  lateral  pouches  of  the  uterus  in  this 
species  or  in  any  species  of  Ophiotaenia  as  Smith  figures  them.  The 
uterus  which  he  figures  is  characteristic  of  Taenia,  not  of  a  Proteoceph- 
alid.  Unfortunately  the  writer  has  not  been  able  to  examine  a  proglot- 
tid from  the  material  received  from  Smith  that  was  in  the  same  state 
of  development  as  that  shown  in  his  figure.  The  writer  has,  however, 
examined  younger  proglottids  and  these  in  every  case  and  in  every 
essential  particular  agreed  with  the  specimens  received  from  other 
sources. 

There  can  be  no  doubt  that  Smith's  specimens  belong  to  the  species 
Crepiddhothrium  gerrardii  Monticelli,  hence  the  name  Taenia  eunectes 
is  a  synonym  of  C.  gerrardii  and  should  be  suppressed. 

This  is  the  only  known  species  in  the  genus  Crepidobothriura,  It 
finds  its  nearest  congeners  among  the  larger  species  of  Ophiotaenia. 
While  in  the  structure  of  the  proglottids  and  in  the  arrangement  of  the 
genital  organs  this  species  agrees  almost  perfectly  with  the  Ophiotaenia 


259]  PROTEOCEPHALIDAE—LA  RUE  259 

there  remain  two  characters  which  are  deemed  of  sufficient  value  to 
warrant  a  separation  of  the  snake  Proteoeephalids  into  two  genera. 
These  characters  are  the  structure  of  the  suckers  and  the  length  of  the 
neck.  The  first  character  has  already  been  discussed.  All  the  known 
species  of  Ophiotaenia  have  relatively  long  necks,  i.  e.,  2-8  mm.  long 
while  C  gerrardii  has  a  short  neck.  Monticelli  was  amply  justified  in 
separating  this  species  from  his  genus  Tetracotylus,  sensu  latu.  How- 
ever since  it  has  been  shown  that  his  type  species  of  Tetracotylus  should 
be  separated  from  the  genus  Proteocephalus  and  even  from  the  family 
Proteocephalidae,  his  arguments  in  justification  of  his  action  are  not 
repeated. 

Monticelli 's  (1899)  diagnosis  of  the  genus  Crepidobothrium  reads: 

"Capo  inerme,  grande,  rigonfio,  piramidato :  ventose  ellittiche,  posteriormente 
a  margine  interrotto  e  rientrante  nel  cavo  della  ventosa.  Aperture  genitali  margi- 
nali,  irregolarmente  alternanti.  Orifizio  dell'antro  genitale  subventrale,  largo : 
sbocco  della  vagina  dorsalmente  e  disotto  alia  tasca  del  pene :  ovario  piccolo : 
testicoli  piccolissimi,  numerosi,  coUocati  nel  mezzo  della  proglottide,  internamente 
alia  musculatura  longitudinale  interna,  in  unica  serie.  Utero  allungato,  tubolare, 
sacciforme.  Vitellogeni  piccoli,  numerosi,  periferici.  Uova  ravvolte  in  una  capsula 
gelatinosa." 

His  diagnosis  does  not  require  much  amplification  but  may  be  re- 
stated thus.  Crepidohothrium  Monticelli:  Head  large,  swollen,  pyra- 
midal, tetragonal,  unarmed.  Fifth  sucker  vestigial.  Suckers  inversely 
cordate,  posterior  margin  interrupted  and  re-entrant  into  sucker  cavity. 
Genital  aperture  marginal,  irregularly  alternating.  Vaginal  opening 
dorsal  to  cirrus-pouch.  Vagina  anterior  or  posterior  to  cirrus-pouch. 
Testes  in  two  lateral  fields  anterior  to  the  ovary.  Ovary  bilobed,  lobes 
irregular,  relatively  small.  Uterus  tubular,  median,  possessing  in  ripe 
proglottids  numerous  lateral  outpocketings.  All  genital  organs  within 
inner  longitudinal  musculature  except  portion  of  cirrus-pouch  and 
vagina.  Eggs  small,  provided  with  three  (?)  membranes,  outer  one 
gelatinous. 

Type  of  genus: — Crepidohothrium  gerrardii  (Baird)  from  Boa  con- 
strictor L.  (type  host)  ;  So.  America  (Bra2dl)   (type  locality). 

In  the  following  table  (pages  260-267)  the  principal  data  relating 
to  the  species  of  Ophiotaenia  and  Crepidobothrium  are  brought  together : 


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268  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [268 


OOCHORISTICA  CRYPTOBOTHRIUM  (von  Linstow)  La  Rue 

1906 :    Ichthyotaenia  crypt  oh  othrium     von  Linstow     1906 :185 
1911 :     Odchoristica  crypt  oh  othria  La  Rue  1911 :481 

^  Under  the  name  of  Ichthyotaenia  cryptohothrium  von  Linstow 
(1906)  described  a  cestode  from  the  intestine  of  a  tree-snake,  Chrysope- 
lea  ornata  Russell,  Kurunegala.  La  Rue  (1911)  stated,  but  without 
assigning  reasons  therefor,  that  this  species  did  not  belong  in  the  genus 
Ichthyotaenia  or  Proteocephalus  but  in  the  genus  Ooehoristica  Liihe. 

A  careful  examination  of  von  Linstow 's  description  and  figures 
shows  that  this  species  cannot  belong  to  the  Proteocephalidae.  The 
position  of  the  vitellaria  in  the  Proteocephalidae  as,  indeed,  in  the  whole 
order  of  the  Tetraphyllidea  is  lateral,  the  vitellaria  extending  nearly 
the  full  length  of  the  proglottid.  In  this  species  the  vitellaria  are 
globular  masses  lying  just  posterior  to  the  ovary.  This  relation  occurs 
in  the  Cyclophyllidea.  Species  of  Odchoristica  Liihe  show  this  relation 
and  since  this  genus  includes  other  species  of  cestodes  parasitic  in  snakes 
it  seems  probable  that  von  Linstow 's  species  belongs  here.  La  Rue 
(1911)  therefore  proposed  for  it  the  specific  name  Odchoristica  crypto- 
hothria  (von  Linstow)  La  Rue.  The  specific  name  should  end  in 
"ium"  not  "ia"  as  he  made  it. 

Unless  the  invaginated  or  infolded  condition  of  the  head  is  a  con- 
stant feature  it  can  have  neither  generic  nor  specific  importance.  Ap- 
parently not  enough  is  known  concerning  the  heads  of  this  species  to 
establish  the  value  of  this  condition  as  a  diagnostic  character. 


MONTICELLIA  CORYPHICEPHALA  (Monticelli)  La  Rue 
[Figs.  94-96,  132,  181,  186] 


1891 
1894 
1896 
1900 
1911 


Tetracotylus  coryphicephala  Monticelli  1891 :151-174 

Ichthyotaenia  coryphicephala  Lonnberg  1894:803 

Ichthyotaenia  coryphicephala  Riggenbach  1896:267 

Ichthyotaenia  coryphicephala  Braun  1894-1900:1680-1681 

Monticellia  coryphicephala  La  Rue  1911:474 


Specific  Diagnosis:    Characters  of  genus.    Head  dilated,  somewhat 
triangular  or  globose,  when  seen  from  the  front  swollen,  sub-quadrate, 


269]  PROTEOCEPHALIDAE—LA  RUE  269 

anteriorly  elongated  to  form  a  conical  elevation,  unarmed  and  not  very 
prominent,  distinct  from  neck.  Breadth  of  head  perhaps  0.35  mm. 
Suckers  four,  fairly  prominent,  oval  in  shape,  about  0.15  mm.  long, 
situated  at  four  angles  of  head.  No  fifth  sucker.  Neck  very  long,  nar- 
row. First  proglottids  short,  middle  ones  almost  rectangular,  posterior 
and  last  ones  decidedly  rectangular,  swollen  in  the  middle.  Ripe  pro- 
glottids about  1.80-2.0  mm.  broad  by  0.68-1.0  mm.  long  by  about  0.34 
mm.  thick.  Proglottids  attached  by  full  width.  Intersegmental  furrows 
shallow.  Corners  of  proglottids  not  marked.  Segmentation  indistinct. 
Genital  organs  as  in  genus.  Genital  pore  marginal,  irregularly 
alternating  in  first  one-fourth  or  one-fifth  of  proglottid  length.  No  geni- 
tal papilla.  Testes  spheroidal,  0.05-0.085  mm.  in  maximum  dimension, 
numerous,  100  or  more,  situated  in  single  dorsal  field  outside  of  inner 
longitudinal  muscle  sheath.  Vas  deferens  a  mass  of  coils  inside  the  inner 
muscle  sheath.  Cirrus-pouch  0.286-0.315  mm.  long  by  0.094-0.105  mm. 
broad.  Length  of  cirrus-pouch  5  to  6  to  6^/^  times  into  proglottid 
breadth.  Ductus  ejaculatorius  describing  numerous  coils.  Cirrus  slen- 
der, unarmed.  Vagina  and  vaginal  opening  always  anterior  to  cirrus- 
pouch.  Vagina  never  crossing  latter.  Sphincter  vaginae  and  receptacu- 
lum  seminis  present.  Vagina  within  inner  muscle  sheath.  Vitellaria 
lateral, .  extensive,  ventral,  follicles  forming  a  single  layer  in  cortical 
parenchyma.  Ovary  posterior,  bilobed,  augmented  by  large  protuber- 
ances, projecting  dorsally,  partially  outside  of  inner  muscle  sheath. 
Organs  of  interovarial  space  as  in  Proteocephalidae.  Uterus  with  many 
long  slender  lateral  outpocketings,  ventral,  and  outside  of  inner  muscle- 
sheath.  Egg-membranes  not  observed.  Embryos  0.015-0.017  mm.  in 
diameter. 

Habitat :    Intestine  of  Silurus  sp.,  locality  not  stated. 
Type :    Slides  in  collection  of  Monticelli,  from  specimen  No.  571  in 
the  Zoological  Museum  at  Vienna. 

This  species  was  first  described  from  Silurus  sp.  by  Monticelli 
(1891:151-174).  For  this  and  nineteen  other  species  of  cestodes  from 
fish  he  proposed  the  genus  Tetracotylus,  without,  however,  naming  a 
type  species.  That  genus  was  based  on  Taenia  coryphicephala,  the  spe- 
cies which  he  described  most  completely.  Lonnberg  (1894:803)  listed 
this  as  one  of  the  species  of  Ichthyotaenia  without  entering  into  a  dis- 
cussion of  its  anatomy.  Riggenbach  (1896:267)  listed  this  among  the 
species  of  Ichthyotaenia.  Braun  (1894-1900:1680-1681)  states  that  Mon- 
ticelli 's  genus  Tetracotylus  is  based  on  the  species  Taenia  coryphicephala 
but  that  the  generic  name  suggested  by  Monticelli  is  a  homonym  of  the 
name  Tetracotyle  suggested  by  Filippi  (1854).    He  said  moreover  that 


270  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [270 

Tetracotylus  had  been  used  in  place  of  Tetracotyle.  For  these  reasons 
the  name  Ichthyotaenia  should  be  used  in  place  of  it. 

La  Rue  (1911:474)  established  for  this  and  certain  other  species  the 
genus  Monticellia  and  the  family  Monticellidae. 

This  study  is  based  upon  data  secured  from  Monticelli's  (1891) 
paper  and  also  upon  Monticelli's  type  slides  which  Professor  H.  B. 
"\yard  very  kindly  secured  for  study.  This  material  originally  came 
from  the  zoological  museum  at  Vienna.  The  host  of  the  species  is  Silu- 
rus  sp. 

Monticelli  (1891)  is  quoted  on  the  external  characters  for  the 
writer  has  examined  only  prepared  slides  of  this  species : 

"Capo  slargato  subtriangolare,  visto  di  fronte,  rigonfiato,  subquadrato,  ante- 
riormente  allunganto  a  formare  un  cocuzzolo  conico,  inertne,  non  molto  proeminente, 
ben  distinto  dal  collo.  Le  quattro  ventose  mediocri  non  molto  proeminenti,  disposte 
ai  quattro  angoli  del  capo.  Collo  lunghissimo,  ristretto.  Prime  proglottidi  brevi, 
mediane  subrettangolari,  posteriori  ed  ultime  decisamente  rettangolari,  rigonfie  nel 
mezzo.     Aperture  genitali  marginal!  irregolarmente  alternanti." 

Monticelli  (1891)  gives  no  dimensions  of  the  head,  neck  and  strobila. 

One  of  Monticelli's  slides  showed  the  head  of  Tetracotylus  coryphi- 
cephala  in  toto.  This  head  which  had  been  badly  compressed  and 
somewhat  broken  by  that  treatment  measured  0.459  mm.  broad.  This 
dimension,  however,  is  too  great  and  it  seems  that  the  normal  head 
would  not  measure  over  0.35  or  perhaps  0.375  mm.  The  head  should  be 
described  as  somewhat  globose  rather  than  sub-triangular.  The  suckers 
could  not  be  measured  except  in  length  which  was  about  0.150  mm. 
Monticelli's  drawing  of  the' head  (Fig.  132)  shows  that  the  suckers  are 
irregularly  oval  in  shape,  the  longitudinal  axis  of  the  suckers  being  the 
greater.  The  neck  in  this  slide  was  too  badly  crushed  to  permit  accurate 
measurement.  It  was  cut  off  3  or  4  mm.  from  the  head  and  showed  no 
traces  of  segmentation.  No  young  proglottids  were  to  be  found  in  the 
slides  examined  by  the  writer.  Four  representative  ripe  proglottids 
were  measured.  Their  dimensions  were  1.95  mm.  broad  by  0.68  mm. 
long,  1.85  by  0.75  mm.,  2.0  by  0.75  mm.,  1.80  by  1.0  mm.  Transections 
of  ripe  proglottids  are  about  0.34  mm,  thick.  The  proglottids  are  at- 
tached by  their  full  width.  The  angles  of  the  segments  can  scarcely  be 
seen  and  the  intersegmental  furrows  are  very  shallow.  These  observa- 
tions were  made  on  a  limited  amount  of  material  so  it  is  possible  that 
some  of  the  statements  just  made  would  require  modification  if  consid- 
erable good  material  could  be  examined. 

The  genital  pore  (Figs.  95,  186)  is  marginal,  irregularly  alternat- 
ing and  situated  at  the  end  of  the  first  one-fifth  to  one-fourth  of  the 


271]  PROTEOCEPHALIDAE—LA  RUE  271 

proglottid.  There  is  no  genital  papilla.  The  genital  pore  (Fig.  181) 
leads  into  a  small  genital  atrium  into  which  both  cirrus-pouch  and 
vagina  open.  The  vagina  always  lies  anterior  to  the  cirrus-pouch  and 
its  opening  is  also  clearly  anterior  to  the  opening  of  the  male  ducts. 
Monticelli  notes  a  sort  of  sphincter  muscle  about  the  genital  atrium 
(Fig.  181).  He  states  that  this  sphincter  draws  the  opening  of  the 
vagina  and  cirrus-pouch  very  closely  together. 

The  testes  are  spheroidal.  They  measure  from  0.05  by  0.05  to  0.05 
by  0.06  to  0.07  by  0.085  mm.,  number  about  100  or  even  more,  and  are 
situated  (Fig.  95)  dorsally  in  a  field  which  covers  the  entire  proglottid 
except  that  region  occupied  by  the  ovary  and  by  a  narrow  strip  on  either 
lateral  margin  of  the  segment.  There  is  no  free  median  zone  as  Monti- 
celli states  and  as  he  shows  in  his  figure,  reproduced  (Fig.  186).  His 
statement  that  they  are  not  numerous  is  incorrect.  The  testes  (Fig,  94) 
are  outside  of  the  inner  longitudinal  muscle  layer,  a  condition  which 
does  not  occur  in  any  species  of  the  Proteocephalidae  nor,  so  far  as  can 
be  ascertained  by  the  writer,  in  any  family  of  Tetraphyllidea.  A  careful 
study  of  Braun's  (1894-1900)  great  monograph  on  the  cestodes  gives  no 
reason  for  believing  that  this  condition  has  been  observed  heretofore  in 
any  order  of  cestodes.  In  the  Pseudophyllidea  he  states  definitely  that 
the  testes  occur  in  the  medullary  layer.  He  makes  the  same  statement 
for  the  Tetraphyllidea  and  the  Cyclophyllidea.  And  as  for  the  genital 
organs  in  the  Diphyllidea  and  the  Trypanorhyncha  he  states  that  they 
are  as  in  the  Tetraphyllidea.  The  location  of  the  testes  in  this  species 
then  is  unique. 

The  vasa  efferentia  have  not  been  observed.  The  vas  deferens  forms 
an  elongated  mass  of  coils  which  extends  from  the  cirrus-pouch  toward, 
but  not  reaching,  the  mid-field.  This  mass  of  coils  lies  just  a  little  poste- 
rior to  the  cirrus-pouch  and  within  the  layer  of  inner  longitudinal  mus- 
cles. The  duct  of  the  vas  deferens  is  not  of  uniform  diameter  through- 
out but  is  narrow  in  its  beginning  region  then  dilated  and  again  con- 
stricted just  before  entering  the  cirrus-pouch.  With  its  swollen  coils  the 
vas  deferens  functions  as  a  vesicula  seminalis.  The  cirrus-pouch  (Fig. 
181)  is  somewhat  muscular,  its  muscles  best  developed  near  the  outer 
end.  Its  length  varies  from  0.286  to  0.315  and  even  to  0.340  mm.  and 
its  breadth  from  0.094  mm.  in  the  longer  pouches  to  0.105  mm.  in  the 
shorter  ones.  The  pouch  tapers  frequently  to  a  blunt  point  at  the 
inner  end  and  is  usually  broadest  near  its  opening.  Its  length  goes  5  to 
6  or  6^  times  into  the  proglottid  breadth.  The  ductus  ejaculatorius 
(Fig.  181)  is  slender  and  has  thin  walls.  It  makes  numerous  intricate 
coils  before  passing  over  into  the  cirrus.    The  protruded  cirrus  has  not 


272  ILLIXOIS    BIOLOGICAL    MONOGRAPHS  [272 

been  observed.  As  the  cirrus  appears  in  the  cirrus-pouch  it  is  weakly- 
muscular,  not  large,  and  is  not  armed.  Probably  when  evaginated  it 
would  be  long,  slender,  and  without  a  thick  base. 

As  seen  in  a  toto  preparation  the  female  organs  appear  to  be  ar- 
ranged in  general  as  in  the  Proteocephalidae  but  a  careful  com- 
parison of  this  species  wuth  any  Proteocephalid  reveals  striking  differ- 
ences. The  vagina  (Fig.  95)  always  lies  anterior  to  the  cirrus-pouch 
and  it  never  crosses  the  latter  nor  does  it  cross  the  mass  of  coils  of  vas 
deferens  except  in  rare  cases  and  then  it  crosses  on  the  ventral  side  of 
only  the  most  anterior  coils,  Monticelli  states  that  it  crosses  the  coils 
of  vas  deferens  dorsally.  The  opening  of  the  vagina  is  always  clearly 
anterior  to  the  opening  of  the  cirrus-pouch.  In  a  number  of  species  of 
Proteocephalus  and  also  of  Ophiotaenia  the  opening  is  dorsal  to  the 
cirrus-pouch,  and  the  writer  is  inclined  to  believe  that  if  the  remaining 
species  of  these  genera  were  carefully  investigated  with  that  point  in 
view  many  of  these  too  would  show  the  opening  of  the  vagina  dorsal  to 
the  cirrus-pouch.  Near  the  vaginal  opening  is  a  weak  sphincter  vaginae. 
Beyond  the  sphincter  the  lumen  of  the  vagina  dilates  slightly  for  a  dis- 
tance, perhaps  0.3  mm.,  then  it  is  constricted.  After  arriving  in  the 
mid-field  of  the  proglottid  the  vagina  again  dilates  considerably.  This 
dilatation  persists  through  several  curves  and  coils  until  the  vagina  is 
just  about  to  enter  the  interovarial  space  when  the  vagina  is  greatly 
constricted.  The  dilated  region  is  the  receptaculum  seminis  which  is 
much  more  marked  than  in  any  species  of  Proteocephalus  observed  by  the 
writer.  As  shown  in  transections  (Fig.  94)  the  vagina,  except  its  be- 
ginning portion,  lies  within  the  internal  layer  of  longitudinal  muscles. 
In  this  respect  Monticellia  coryphicephala  agrees  with  the  Proteocepha- 
lidae. 

The  ovary  (Fig.  95)  is  posterior  and  bilobed.  A  narrow,  thin  raid- 
piece  connects  the  lobes  which  are  long  and  broad.  From  the  dorsal 
surface  of  the  distal  region  of  each  lobe  rounded  prominences  arise  and 
extend  toward  the  dorsal  surface  of  the  proglottid.  Many  of  these 
prominences  when  observed  in  toto  preparations  seem  to  have  no  con- 
nection with  the  ovary  and  are  readily  mistaken  for  testes  but  when 
studied  in  transections  they  are  seen  to  be  a  part  of  it.  ]\Ionticelli 's 
drawing  which  is  reproduced  (Fig.  186)  does  not  truly  represent  this 
appearance.  These  dorsal  prominences  and  the  greater  part  of  the 
ovarian  lobes  (Fig.  96)  from  which  they  arise  lie  in  the  cortical  paren- 
chyma outside  of  the  internal  layer  of  longitudinal  muscles.  The  mid- 
piece  and  part  of  the  lobes  of  the  ovary  lie  within  the  muscular  sheath. 
Monticelli  (1891)  failed  to  point  out  this  condition.    Here  is  a  relation. 


273]  PROTEOCEPHALIDAE—LA  RUE  273 

the  like  of  which  the  writer  has  been  unable  to  find  elsewhere  in  all  the 
cestodes.  So  far  as  he  is  able  to  determine  the  ovary  of  other  cestodes 
always  lies  in  the  medullary  parenchyma. 

At  the  posterior  margin  of  the  ovarian  mid-piece  is  situated  the 
oocapt.  This  organ  was  called  the  "sfintere  ovarico"  by  Monticelli. 
The  oocapt  leads  into  the  oviduct  which  after  making  several  coils  re- 
ceives the  vagina.  Then  as  a  fertilization  passage  the  oviduct  continues 
until  it  reaches  the  ootype  which  is  surrounded  by  the  shell  glands. 
Just  before  entering  the  ootype  the  oviduct  receives  the  unpaired  vitel- 
line duct.  The  ootype  discharges  into  the  uterine  passage  which  passes 
ventral  to  the  ovary  and  then  discharges  into  the  uterus.  It  will  be  seen 
from  a  comparison  of  the  above  description  with  drawings  and  descrip- 
tions of  similar  organs  in  the  Proteocephalidae  that  there  is  marked 
agreement  between  Monticellia  and  the  Proteocephalidae  in  these  rela- 
tions. However  this  is  no  more  than  can  be  reasonably  expected  when 
it  is  remembered  that  practically  the  same  plan  for  these  organs  is  fol- 
lowed throughout  the  whole  order  of  Tetraphyllidea  to  which  Monticel- 
lia and  Proteocephalus  belong.  Monticelli  (1891)  takes  occasion  to 
point  out  what  seems  to  be  misinterpretations  on  the  part  of  von  Lin- 
stow  (1891)  in  describing  these  organs  in  Taenia  longicollis.  The  writer 
agrees  with  Monticelli  in  considering  that  von  Linstow  has  mistaken  the 
oocapt  for  the  ootype.  A  farther  discussion  of  this  point  will  be  found 
in  the  description  of  Proteocephalus  longicollis. 

The  vitellaria,  which  in  the  Proteocephalidae  are  lateral,  follicular 
masses,  the  individual  follicles  of  which  are  closely  grouped  about  a 
central  conducting  tubule,  are  in  Monticellia  coryphicephala  (Fig.  95) 
widely  scattered  lateral  follicular  masses  which  lie  in  a  single  ventral 
layer.  In  Proteocephalids  the  vitellaria  are  within  the  inner  longitudi- 
nal muscle  layer  while  in  M.  coryphicephala  they  lie  outside  of  those 
muscles.  The  vitellaria  are  not  only  ventral  but  at  the  margin  of  the 
proglottid  they  turn  up  toward  the  dorsal  surface,  hence  when  viewed 
in  a  toto  preparation  the  vitellaria  at  the  margins  seem  greatly  com- 
pacted. Transections  (Fig.  94)  of  the  proglottids  show  the  method  of 
arrangement  of  the  vitellaria.  The  lateral  longitudinal  vitelline  ducts 
have  not  been  seen  on  account  of  the  poor  histological  condition  of  the 
material.  The  paired  vitelline  ducts  however,  may  be  seen  to  arise  in 
the  lateral  fields  and  to  pass  toward  the  middle  of  the  proglottid  outside 
of  the  longitudinal  muscle  layer.  After  running  for  some  distance  in 
this  relation  the  ducts  turn  dorsally  and  enter  within  the  muscle  layer 
and  then  they  pass  on  to  the  middle  of  the  proglottid  where  the  paired 
vitelline  ducts  unite  to  form  the  unpaired  vitelline  duct  which  discharges 
into  the  ootype.    In  all  the  known  species  of  Proteocephalidae  the  vitel- 


274  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [274 

line  ducts  are  always  within  the  longitudinal  muscle  layer.  Monticelli 
failed  to  note  the  position  of  the  vitelline  ducts  in  M.  coryphicephala. 

The  uterus  (Fig.  95)  in  ripe  proglottids  is  made  up  of  about  18-30 
slender  out-pocketings,  which  extend  well  toward  the  margins  of  the 
proglottids  but  not  as  far  laterad  as  in  certain  species  of  Proteocephalus 
such  as  P.  fallax,  P.  duhius,  and  P.  ambloplitis.  Monticelli  (1891)  states 
that  the  uterus  when  filled  with  eggs  causes  the  ventral  surface  of  the 
proglottid  to  belly  out.  Unfortunately  his  drawing  showing  that  con- 
dition has  not  been  reproduced.  The  uterus  (Fig.  94)  is  ventral  and  it 
too  lies  outside  of  the  internal  layer  of  longitudinal  muscles.  Uterine 
pores  have  not  been  observed.  Monticelli  says  that  they  do  not  exist 
but  that  when  the  uterus  is  full  the  ventral  body  wall  ruptures,  thus 
discharging  the  eggs.  A  careful  study  of  good  material  would  probably 
show  the  presence  of  one  or  more  uterine  pores  such  as  was  found  by 
Kramer  (1892),  Schneider  (1905)  and  by  the  writer  in  a  large  number 
of  Proteocephalids.  Monticelli  gives  no  measurements  of  the  uterine 
eggs.  He  states  that  the  membranes  are  quite  delicate  and  that  the 
uterine  eggs  are  very  small.  In  one  of  Monticelli 's  slides  a  broken  pro- 
glottid rendered  some  of  the  eggs  visible.  The  spheroidal  embryos 
measured  about  0.015  to  0.017  mm.  in  diameter.  The  membranes  had 
become  so  transparent  that  they  could  not  be  measured. 

In  the  above  description  and  in  the  drawings  mentioned  it  has  been 
shown  that  Monticellia  coryphicephala  differs  from  the  Proteocephalidae 
and  especially  from  the  typical  species  of  Proteocephalus  in  a  number 
of  characters  which  concern  the  position  of  the  genital  organs  and  their 
relation  to  the  inner  longitudinal  muscle  layer.  In  all  the  Proteocepha- 
lids which  have  been  sufficiently  investigated  to  determine  these  points 
the  entire  genital  apparatus  with  the  exception  of  a  portion  of  the 
cirrus-pouch  and  vagina  lies  within  the  inner  longitudinal  muscle  layer. 
Moreover,  the  follicles  of  the  vitellaria  are  closely  packed  about  a  central 
duct.  In  Monticellia  the  vitellaria  are  in  two  broad  lateral  fields  the 
follicles  of  which  are  scattered  in  a  broad  single  layer.  The  paired 
vitelline  ducts  for  a  considerable  distance  lie  outside  the  inner  muscle 
layer.  The  testes  and  uterus  are  entirely  outside  this  muscle  layer  and 
the  ovary  is  partly  outside  of  it.  As  in  the  Proteocephalidae  the  vagina, 
vas  deferens  and  part  of  the  cirrus-pouch  are  within  the  inner  muscle 
layer.  On  account  of  these  marked  differences  between  the  Proteo- 
cephalidae and  the  form  which  Monticelli  named  Tetracotylus  coryphi- 
cephala the  writer  (1911:474)  has  seen  fit  to  establish  a  new  genus 
Monticellia  in  honor  of  Professor  Monticelli. 

The  status  of  Monticelli 's  generic  name  Tetracotylus  has  already 
been  reviewed  in  the  introductory  section  where  the  new  genus  has  been 


275]  PROTEOCEPHALIDAE—LA  RUE  275 

defined  as  has  also  the  new  family  Montieellidae.  In  this  genus  and 
family  probably  belong  Tetracotylus  diesingii  Monticelli,  T.  macrocoty- 
lea  Monticelli  and  Taenia  malopteruri  Fritsch.  These  three  species  are 
all  parasitic  in  the  Siluridae.  Perhaps  Taenia  osculata  Goeze  belongs 
here  also.  It  seems  preferable,  however,  for  the  present  to  consider  this 
last  as  a  species  inquirenda  in  the  genus  Proteocephalus  in  which  it  has 
long  been  considered  to  belong.  More  careful  study  into  the  morphology 
of  Taenia  osculata  is  desirable  in  order  that  its  systematic  position  may 
be  determined. 

MONTICELLIA  MACROCOTYLEA  (Monticelli) 
[Fig.  125] 
Tetracotylus  macrocotylea        Monticelli         1891 


1891 
1896 
1911 


Ichthyotaenia  macrocotylea      Riggenbach      1896:267 
Monticellia  macrocotylea  La  Rue  1911 :474 


This  species  was  described  by  Monticelli  (1891).  Lack  of  sufficient 
good  material  prevented  a  careful  study  of  it.  Riggenbach  (1896:267) 
listed  this  form  in  the  genus  Ichthyotaenia.  La  Rue  (1911:474)  in- 
cluded this  species  in  the  genus  Monticellia.  Monticelli 's  original  de- 
scription is  here  quoted:  ''TAENIA  MACROCOTYLEA.— Questa 
Taenia,  che  indico  cosi  dalle  sue  proeminenti  ventose,  ha  raolte  rassomigli- 
anze  con  le  due  seguenti,  ma  da  entrambe  si  distingue  per  la  caratteris- 
tica  forma  del  capo  che  e  nettamente  distinto  dal  collo  e  per  il  eoUo  piu 
breve.  Le  aperture  genitali  sono  marginali  irregolarmente  alternanti. 
Lo  stato  deir  individuo  che  posseggo  non  mi  ha  permesso  uno  studio  piu 
minuzioso  di  questa  Taenia.     •     •     •     *     * 

Habitat:  intestino  del  SUurus  megacephalus." 

Professor  H.  B.  Ward  very  kindly  secured  Monticelli 's  type  slides 
of  this  species  for  examination.  As  a  result  of  this  examination  the 
writer  is  able  to  add  the  following  data  to  the  original  description. 

The  head  of  this  specimen  is  slightly  crushed.  In  this  condition  it 
measures  1.10  mm.  broad,  a  figure  which  is  probably  too  large  by  0.10- 
0.20  mm.  Its  length  is  about  0.90  mm.  The  suckers  are  prominent  and 
heavily  muscled.  They  measure  0.45  to  0.50  mm.  in  diameter.  The 
sucker  cavity  could  not  be  measured.  The  neck  is  about  0.50  mm.  broad 
just  back  of  the  head.  Its  length  could  not  be  ascertained  for  it  was 
broken  off.  Mature  proglottids  measure  about  0.75  mm.  square.  No 
young  or  ripe  proglottids  could  be  seen.    The  segmentation  is  distinct. 

The  genital  pore  is  marginal,  irregularly  alternating,  and  situated 
in  the  first  one-fourth  of  the  proglottid.    It  is  rendered  prominent  by  a 


276  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [276 

small  genital  papilla.  The  length  of  the  cirrus-pouch  and  the  relations 
of  the  latter  with  the  vagina  could  not  be  ascertained.  The  testes  num- 
ber about  100  or  more,  and  they  are  situated  in  a  single  field  between 
the  vitellaria.  The  vitellaria  are  lateral  follicular  organs,  the  follicles 
of  which  are  scattered  much  as  they  are  in  Monticellia  coryphicephala. 
The  ovary  is  bilobed,  and  it  occupies  a  posterior  position  on  the  pro- 
glottid. The  material,  however,  is  too  poor  to  permit  of  a  careful  study 
of  this  organ.    The  uterus  could  not  be  seen. 

This  species  belongs  in  the  genus  Monticellia  La  Rue.  It  is  differ- 
entiated from  the  other  species  of  the  genus  by  reason  of  its  larger  head 
and  suckers,  its  prominent  genital  pore  and  its  evident  segmentation. 
Unfortunately  the  material  is  so  poor  that  a  careful  study  is  not  per- 
mitted and  as  a  consequence  its  exact  relation  to  the  other  species  of 
the  genus  cannot  be  determined. 


MONTICELLIA  DIESINGII  (Monticelli) 
[Fig.  157] 


1891 
1896 
1911 


Tetracotylus  diesingii  Monticelli  1891 

Ichthyotaenia  diesingii         Riggenbach        1896:267 
Monticellia  diesingii  La  Rue  1911:474 


This  species  was  very  inadequately  described  by  Monticelli  (1891) 
whose  material  was  too  limited  and  too  poor  to  permit  a  careful  stul!y. 
Riggenbach  (1896:267)  listed  it  as  one  of  the  species  of  Ichthyotaenia 
but  gave  no  description  of  it.  La  Rue  (1911:474)  included  this  form 
among  the  species  of  Monticellia.  Monticelli 's  original  description  is 
here  quoted:  "TAENIA  DIESINGII.— Capo  subgloboso  anterior- 
mente  rotendato  appena  distinto  dal  coUo.  Le  quattro  ventose  grandi, 
proeminente  e  molto  muscolari.  CoUo  assai  lungo:  prime  proglottidi 
brevissime  appiattite,  proglottidi  mediane  alquanto  rigonfie  all'  aspetto 
cerciniformi,  posteriori  rettangolari,  ultime  subquadrate.  Aperture  gen- 
itali  irregolarmente  alternanti.  L'apparato  genitale,  da  quanto  mi  e 
riuscito  vedere,    si  assomiglia  a  quello  della  T.  coryphicephala. 

Habitat:  intestino  del  SUurus  dargado." 

Thanks  to  Professor  H.  B.  Ward  the  writer  has  been  able  to  exam- 
ine Monticelli 's  type  slides  of  this  species  and  can  add  the  following 
data  to  the  original  description. 

The  head  is  about  0.30  mm.  in  breadth.  Suckers  measure  about 
0.17  mm.  in  their  maximum  dimension  while  the  opening  of  the  same  is 
about  0.08  mm.  long.     The  musculature  of  the  sucker  is  thick.    Altho 


277]  PROTEOCEPHALIDAE—LA  RUE  277 

the  neck  of  this  specimen  is  broken  into  several  pieces,  its  pieces  total 
about  3-5  mm.  in  length.  The  young  proglottids  are  about  0.05  mm. 
long  by  0.34  mm.  broad.  Fully  mature  proglottids  are  about  0.80  mm. 
broad  by  0.45-0.50  mm.  long.  Since  the  proglottids  are  attached  by  their 
full  width  the  segmentation  is  not  distinct. 

The  genital  pore  is  marginal,  situated  within  the  first  one-third  or 
one-fourth  of  the  proglottid.  Its  position  alternates  irregularly.  The 
cirrus-pouch  is  about  0.265  mm.  long  by  0.105  mm.  broad.  The  ductus 
ejaculatorius  forms  numerous  coils  within  the  cirrus-pouch.  The  cirrus 
is  slender  and  not  strongly  muscular.  It  was  not  seen  protruded.  Testes 
numbering  about  100  are  situated  in  the  whole  dorsal  field  between  the 
vitellaria.  The  coils  of  vas  deferens  could  not  be  seen.  The  vitellaria 
are  lateral  and  the  follicles  seem  to  be  scattered  much  as  in  Monticellia 
coryphicephala.  It  was  not  difficult  to  determine  that  the  bilobed  ovary 
is  situated  in  the  posterior  part  of  the  proglottid  but  it  was  impossible 
to  determine  its  true  character.  The  vagina  lies  anterior  or  posterior 
(?)  to  the  cirrus-pouch.  Uterus  and  eggs  could  not  be  observed  in  this 
material. 

It  is  unfortunate  that  it  is  impossible  to  give  a  more  complete 
description  than  is  here  given.  The  specimens  were  stained  with  car- 
mine which  at  its  best,  does  not  yield  good  outlines  of  organs  in  the 
cestodes.  In  this  case  the  differentiation  was  poor  making  a  careful 
study  out  of  the  question.  Altho  much  to  be  desired  it  did  not  seem 
advisable  on  account  of  the  extremely  fragile  condition  of  the  specimens 
to  attempt  a  restaining  of  the  material,  even  had  Professor  Monticelli 
been  willing  that  the  attempt  be  made. 

The  study  of  this  material  has  convinced  the  writer  that  this  species 
belongs  to  the  genus  Monticellia  La  Rue.  It  is  closely  related  to  M.  cory- 
phicephala but  seems  to  be  quite  distinct  from  that  species.  Without 
an  examination  of  more  material  many  points  must  remain  in  doubt. 

MONTICELLIA  MALOPTERURI  (Fritsch)  La  Rue 
[Figs.  154,  166] 

1886:  Taenia  malopteruri  Fritsch  1886:103-108 

1891 :  Taenia  malopteruri  Monticelli  1891 

1896:  Ichthyotaenia  malopteruri  Riggenbach  1896:267 

1911 :  Monticellia  malopteruri  La  Rue  1911 :474 

Specific  Diagnosis:  Characters  of  genus.  Length  unknown. 
Breadth  as  much  as  1.5  mm.  Scolex  fairly  large,  0.45  mm.  in  breadth, 
quadrangular,  bearing  four  prominent  suckers.     Rostellum   ( ?)   hemi- 


27S  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [278 

spherical,  armed  with  many  small  hooks.  Suckers  large,  heavily  muscled, 
protuberant,  0.25  mm.  in  diameter.  Neck  short,  nearly  broad  as  head. 
Mature  proglottids  nearly  quadrate.  Ripe  proglottids  broader  than 
long,  1.46  mm.  broad  by  1.1  mm.  long.  Genital  organs  as  in  genus. 
Genital  pore  marginal,  irregularly  alternating,  near  middle  of  pro- 
glottid. Pore  large,  marked  by  a  thickening  of  margin.  Cirrus,  when 
protruded,  short  and  filiform.  Cirrus-pouch  short  and  relatively  small, 
extending  about  one-fifth  across  proglottid  breadth.  No  coils  of  ductus 
ejaculatorius  in  pouch.  Vas  deferens,  a  mass  of  coils  extending  to  mid- 
dle of  proglottid.  Testes  filling  up  field  between  vitellaria,  about  120  in 
number.  Vagina  posterior  to  cirrus-pouch.  No  coils  of  vagina  anterior 
to  ovary.  Ovary  a  bilobed,  winglike  structure.  Vitellaria  follicular, 
situated  in  lateral  fields.  Uterus,  a  median  tube  with  18-20  lateral 
pouches.    Eggs,  not  described. 

Habitat:    In  the  intestine  of  Malopterurus  electricus  ( ?),  Egypt. 

Fritsch  described  and  figured  this  form  in  1886  as  Taenia  malop- 
teruri.  Monticelli  (1891)  included  this  species  in  his  group  of  fish 
cestodes.  He  did  not  describe  it.  Riggenbach  (1896)  considered  this  a 
species  of  Ichthyotaenia.  La  Rue  (1911:474)  considered  this  to  be  a 
species  of  Monticellia. 

Fritsch 's  diagnosis  reads:  ^'Taenia  malopteruri,  Caput  quadrangu- 
lare,  acetabulis  quattuor,  angulariter  positis  robustis.  Rostellum  hemis- 
phaericum,  spinis  obtusis  vel  tuberculis  minimus  ornatum.  Collum 
mediocre.  Aperturae  genitalium  marginales  leviter  circumvallatae. 
Corpus  sulco  profundo  per  medium  impressum.  Articuli  adulti  dimido 
fere  longiores  quam  lati,  proglottides  breves,  contracti.  Habitat  in 
intestino  Malopteruri."  The  description  given  by  Fritsch  is  very  in- 
complete. However,  he  figures  the  head  and  a  nearly  ripe  proglottid, 
stating  the  magnification  of  the  same.  Since  this  is  true  one  can  secure 
some  size  relations  by  measuring  the  organs  portrayed  in  the  drawings. 
In  doing  this  care  must  be  used  because  in  some  respects  the  drawings 
are  not  altogether  clear,  nor  can  one  always  determine  the  character  of 
certain  organs.  Nevertheless  the  writer  has  used  these  measurements  in 
the  full  realization  that  they  were  inaccurate  but  believing  that  they 
probably  are  correct  within  the  limits  of  variation  of  the  species.  Besides 
the  study  of  these  drawings  the  description  is  based  upon  Fritsch 's 
(1886)  description. 

The  length  is  not  stated.  In  breadth  the  worm  may  measure  as 
much  as  1.5  mm.  The  scolex  (Fig.  154)  is  of  fair  size,  0.45  mm.  in 
diameter.  It  is  quadrangular  and  bears  four  prominent  suckers  and  an 
hemispherical  rostellum   (?),  armed  with  many  small  hooks.     Suckers 


279]  PROTEOCEPHALIDAE—LA  RUE  279 

are  large,  0.25  mm.  in  diameter,  heavily  muscled,  and  protuberant  from 
the  angles  of  the  head.  The  neck  is  short  but  nearly  as  broad  as  the 
head.  Fritsch  failed  to  state  the  proportions  of  the  young  proglottids, 
nor  are  the  latter  figured.  Mature  proglottids  are  nearly  quadrate  while 
ripe  ones  are  somewhat  broader  than  long.  Ripe  proglottids  measure 
about  1.46  mm.  broad  by  1.1  mm.  long.  The  genital  organs  are  arranged 
according  to  the  type  of  the  genus.  The  marginal  genital  pore  alter- 
nates from  side  to  side.  It  is  situated  near  the  middle  of  the  proglottid, 
is  of  large  size  and  is  marked  by  a  considerable  thickening  of  the  mar- 
gin. The  cirrus  is  sometimes  protruded  and  is  then  short  and  slender. 
The  cirrus-pouch  is  relatively  small  and  short.  Its  length  goes  into  the 
proglottid  breadth  about  five  times.  The  ductus  ejaculatorius  forms  no 
coils  within  the  cirrus-pouch.  The  coils  of  the  vas  deferens  form  a  knot 
extending  from  the  cirrus-pouch  to  the  middle  of  the  proglottid.  Testes 
are  irregularly  scattered  between  the  vitellaria  and  the  ovary.  Fritsch 
figures  about  120  of  them.  His  drawing  which  is  reproduced  here  (Fig. 
166)  does  not  show  them  in  the  mid-field.  This  may  be  explained  in 
three  ways:  first,  that  the  testes  occupy  only  the  lateral  fields  of  the 
proglottid,  second,  that  the  preparation  was  of  such  a  nature  that  the 
testes  if  present  could  not  be  seen  on  account  of  the  eggs  in  the  uterus, 
or  third,  that  they  were  poorly  stained.  Without  having  seen  the 
specimens  on  which  Fritsch 's  description  is  based  or  specimens  known 
to  belong  to  the  same  lot  or  from  the  same  host  species  one  is  unable  to 
judge  on  this  point.  It  is  well,  perhaps,  to  call  attention  to  the  fact 
that  testes  in  ripe  proglottids  do  not  take  the  stain  well  nor  are  they 
easily  seen  through  the  mass  of  eggs  in  the  uterus. 

The  vagina  opens  posterior  to  the  cirrus-pouch.  "Whether  it  ever 
opens  anterior  to  the  pouch  is  not  stated.  There  are  no  coils  of  the 
vagina  anterior  to  the  ovary.  The  ovary  is  bilobed,  each  lobe  being 
somewhat  winglike.  It  is  in  the  posterior  part  of  the  proglottid.  The 
lobes  are  apparently  follicular  at  the  outer  ends,  tho  this  feature  could 
not  be  determined  with  any  accuracy  unless  an  examination  of  Fritsch 's 
specimens  were  made.  The  vitellaria  are  long  follicular  masses  in  either 
lateral  field.  From  the  drawing  the  follicles  of  the  vitellaria  seem  to 
be  compacted  rather  than  widely  scattered  as  in  the  type  of  the  genus. 
The  uterus  is  a  median  tube  from  which,  apparently,  many  lateral 
pouches  arise.  In  the  drawing  by  Fritsch  these  pouches  are  rather 
indistinct,  but  about  18-20  on  either  side  can  be  seen  with  some  degree 
of  certainty.  In  the  drawing,  reproduced  (Fig.  166)  these  structures 
are  made  more  distinct  than  in  the  original.  The  eggs  were  not  de- 
scribed by  Fritsch.  % 


280  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [280 

The  systematic  position  of  this  species  is  not  readily  determined 
from  the  data  at  hand.  Fritsch's  description  leaves  much  to  be  desired 
and  his  figures  are  not  sufficiently  definite  to  render  possible  a  positive 
determination.  Nevertheless,  certain  features  of  the  anatomy  of  the 
worm  are  so  well  delineated  that  some  definite  conclusions  can  be 
reached.  The  rounded  summit  of  the  head  covered  wath  numerous 
spines  and  the  very  prominent  suckers  which  seem  to  be  set  on  the 
surface  of  the  head  are  not  typical  of  Proteocephalus.  Species  of  that 
genus  lack  spines  on  the  head  and  their  suckers  are  never  as  prominent 
as  in  the  species  under  consideration.  The  head  agrees  very  well  with 
that  of  Moniicellia  coryphicephala  Monticelli  in  regard  to  the  suckers. 
The  latter  species  has  no  spines.  In  the  genital  organs  there  seems,  in 
the  main,  to  be  a  good  agreement  with  the  members  of  the  genus  Pro- 
teocephalus. Nevertheless,  a  careful  examination  and  comparison  of 
Fritsch's  figure  of  the  proglottid  (Fig.  166)  with  the  writer's  figure  of 
Monticellia  coryphicephala  (Fig,  95)  shows  a  certain  resemblance  that 
cannot  be  overlooked.  In  both  species  there  is  the  more  or  less  indefi- 
nitely outlined  ovary  made  up  of  follicles  at  the  outer  regions.  These 
follicles  seem  to  be  smaller  in  this  species  than  in  M.  coryphicephala. 
The  vitellaria,  apparently,  are  not  as  extensive  in  the  former  as  in  the 
latter,  yet  they  are  different  from  those  in  Proteocephalus.  The  lips 
of  the  genital  sinus  are  much  more  prominent  in  this  species  than  in 
Proteocephalus.  This  character  was  not  mentioned  by  Monticelli  as 
being  present  in  Tetracotylus  nor  has  the  writer  noted  it  in  his  specimens. 

In  view  of  the  facts  above  presented  the  writer  must  conclude  that 
the  species  does  not  belong  to  the  genus  Proteocephalus  Weinland. 
With  some  reservation  this  species  is  assigned  to  the  genus  Monticellia 
La  Rue.  The  species  is  then  to  be  knoAvn  as  Monticellia  malopteruri 
(Fritsch)  La  Rue.  The  species  of  cestodes  infesting  the  Siluridae  merit 
much  more  careful  study  by  modern  methods.  As  yet  their  structure 
and  relationships  are  almost  unknown.  The  literature  contains  but  few 
references  to  cestodes  of  this  group.  The  writer  has  noted  the  descrip- 
tions of  the  species  Tetracampos  ciliotheca  Wedl  and  Marsypocephalus 
rectangulus  Wedl  reported  by  Wedl  (1861)  from  Heterobranchus  an- 
guUlaris.  The  former  because  of  its  ventral  genital  pore,  ciliated  embryo 
and  two  bothria  evidently  belongs  to  the  order  Pseudophyllidea.  The 
latter  species  may  belong  somewhere  in  the  Proteocephalidae  or  Monti- 
cellidae  but  not  sufficient  data  are  given  to  warrant  a  positive  determi- 
nation. Wedl  (1861)  also  reported  Scolex  syndontis  from  Syndontis 
schal  and  from  Heterobranchus  anguUlaris.  These  specimens,  likewise, 
cannot  be  definitely  placed. 


281]  PROTEOCEPHALIDAE—LA  RUE  281 


DISTRIBUTION  OF  PROTEOCEPHALIDAE 

The  data  upon  which  this  section  is  based  are  meager  yet  it  has 
seemed  desirable  to  work  over  the  available  records  to  find  out  if  there 
be  any  principles  governing  the  distribution  of  the  group,  to  find  out 
the  relation  of  the  parasite  to  the  primary  hosts  and  perhaps  to  the 
secondary  hosts.  This  section  falls  naturally  into  two  parts:  A,  con- 
cerning the  Proteocephalids  infesting  the  amphibians  and  reptiles,  and 
B,  concerning  those  species  which  infest  fish.  The  data  under  the  first 
heading  are  grouped  into  certain  tables  and  a  discussion  of  the  contents 
of  the  tables.  The  data  bearing  on  the  second  heading,  i.  e.,  on  the  fish 
Proteocephalids  are  grouped  under  the  following  rubrics:  List  of  ces- 
tode  species  with  hosts  and  locality  of  finding;  Distribution  according 
to  continents;  Host  species  harboring  two  or  more  Proteocephalid  spe- 
cies, with  the  distribution  of  the  latter;  Proteocephalids  found  in  more 
than  one  host  species,  a  discussion  of  each  case  including  notes  on  the 
distribution  and  food  habits  of  the  hosts;  Families  of  fish  from  which 
Proeocephalids  are  known  and  the  species  of  the  latter. 

Distribution  op  Proteocephalids  of  Amphibia  and  Reptilia 

Amphibian  Proteocephalids  are  known  from  but  two  continents, 
North  America  and  Australia.  These  species  are  Ophiotaenia  filaroides 
and  0.  lonnhergii  for  North  America  and  0.  hylae  for  Australia.  The 
North  American  species  are  from  the  Caudata  while  the  Australian 
species  is  from  one  of  the  Salientia.  Reptilian  Proteocephalids  are 
known  from  all  the  continents.  In  North  America  are  found  Ophiotae- 
nia marenzelleri,  0.  grandis,  0.  perspicua,  and  0.  lactea  sp.  inq.  In 
South  America  are  found  0.  calmettei,  0.  racemosa,  and  Crepidoboth- 
rium  gerrardii.  In  Europe  a  single  species  is  known,  namely  0.  nat- 
tereri.  Africa,  likewise,  has  yielded  but  a  single  species,  0.  punica. 
Asia  including  the  tropical  islands  usually  included  in  the  Oriental 
region  has  three  species,  Acanthotaenia  shipleyi,  Ophiotaenia  pigmen- 
tata,  and  0.  trimeresuri.  In  Australia,  i.  e.,  in  the  Australian  region 
which  includes  new  Guinea,  there  are  no  species  of  Ophiotaenia  from 
reptiles  but  there  are  four  species  of  Acanthotaenia  from  reptiles,  viz. 
A  biroi,  A.  saccifera,  A.  tidswelU,  and  A.  gallardi. 

To  sum  up  this  data  there  are  species  of  Ophiotaenia  from  each 
continent,  species  of  Acanthotaenia  from  two  continents,  Asia  and  Aus- 
tralia, and  one  species  of  Crepidobothrium  from  South  America.     As 


282  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [282 

yet  none  of  these  species  are  known  to  occur  in  more  than  one  conti- 
nent. The  distribution  of  each  species  seems  narrow.  There  are,  how- 
ever, but  few  records  and  the  data  are  incomplete.  When  more  records 
are  available  it  may  be  found  that  some  of  these  species  may  have  a  far 
wider  distribution.  In  all  the  cases  known  the  hosts  are  aquatic  or 
semiaquatic  animals  and  they  feed  in  part  at  least  upon  animals  which 
like  themselves  are  aquatic  or  semiaquatic  in  their  habits.  It  has  not 
been  possible  to  work  over  the  rather  scant  literature  on  the  food  habits 
of  these  animals  hence  no  data  on  that  topic  are  here  presented.  It  is 
to  be  hoped  that  someone  may  make  a  study  of  the  food  of  these  hosts 
and  thus  enable  to  be  made  a  determination  of  the  relationships  of  these 
parasites  to  each  other  and  to  their  hosts. 

An  accompanying  table  gives  the  names  of  the  parasites,  their  hosts 
and  the  locality  of  the  collection.  In  another  table  the  parasites  have 
been  grouped  according  to  the  order  and  family  of  the  host.  The  latter 
table  shows  that  there  are  Proteocephalid  species  from  Caudate  and 
Salientian  Amphibians,  from  the  Varanidae,  a  family  of  Lacertilian 
Reptiles,  and  from  three  families  of  Serpentes  (Ophidia).  These  fami- 
lies are  Boidae,  Colubridae,  and  Viperidae.  A  single  subfamily  in  each 
of  the  Boidae  and  Viperidae  furnish  hosts  for  Proteocephalids  while  in 
the  family  Colubridae  one  subfamily  belonging  to  each  of  the  three  se- 
ries furnishes  host  species.  At  the  bottom  of  the  table  is  placed  a  species 
which  tho  reported  to  have  been  taken  from  a  dog  is  certainly  not  a  dog 
cestode.  Its  nearest  relatives,  anatomically,  are  the  Proteocephalids  of 
snakes  and  particularly  the  Proteocephalids  of  the  Crotalinae.  It  has 
been  suggested  elsewhere  that  the  real  host  was  probably  one  of  the 
Crotalinae.  Because  of  the  small  amount  of  data  it  has  not  seemed 
justifiable  to  draw  many  general  conclusions  in  regard  to  these  para- 
sites. 


283] 


PROTEOCEPHALIDAE—LA  RUE 


283 


Hosts  and  Localities  Reported  for  Proteocephalids  of  Amphibia  and  Reptilia 


Cestode  species 

Hosts 

Locality 

Acanthotaenia   biroi    (von 

Varanus  sp. 

New  Guinea 

Ratz) 

A.  gallardi  Johnston 

Pseudechis  porphyriacus 
Shaw 

New  South  Wales 

P.  australis  Gray 

New  South  Wales 

Notechis  scutatus  Peters 

New  South  Wales 

Denisonia  superba  Gunther 

New  South  Wales 

A.  saccifera   (von  Ratz) 

Varanus  sp. 

New  Guinea 

A.  shipleyi  von  Linstow 

Varanus   (Hydrosaurus) 
salvator 

Ceylon 

A.  tidswelli  Johnston 

Varanus  varius  Shaw 

New  South  Wales 

V.  gouldii  Gray 

Burnett  River,  Australia 

V.  bellii  (a  variety  of  V. 

Burnett  River,  Australia 

varius) 

Crepidobothrium  gerrardii 

Eunectes  murinus  Wagl. 

South  America 

(Baird) 

Boa  constrictor  L. 

South  America 

Ophiotaenia  calmetti  (Bar- 

Lachesis    (Bothrops)    lan- 

Martinique,  West   Indies 

rois) 

ce  olatus  L. 

Sao  Paulo,  Brazil 

0.  filaroides  La  Rue 

Amblystoma  tigrinum 
(Green) 

Nebraska  and  Kansas 

0.  grandis  La  Rue 

Ancistrodon  piscivorus 
Holbr. 

Southern  United  States 

0.  hylae  Johnston 

Hyla  aurea 

New  South  Wales 

0.  lactea  (Leidy)  sp.  inq. 

Matrix  sipedon 

(?)  Eastern  United  States 

0.  lonnbergii  (Fuhrmann) 

Necturus  maculosus 

Ohio  and  Indiana 

0.  marenselleri  (Barrois) 

Ancistrodon  piscivorus 
Holbr. 

(  ?)  Southern  United  States 

O.  nattereri  (Parona) 

Coluber  sp. 

Liguria,  an  Italian  province 

O.  perspictia  La  Rue 

Natrix    rhombifer    Hallo- 
well 

Illinois  and  Oklahoma 

0.  piginentata  (von  Lin- 

Psa mm 0 dynast es    pulver- 

Java  (Semarang) 

stow) 

ulentus  Boie 

0.  punica  (Cholodkovski) 

Dog  (?),  probably  a  snake 

Island  of  Dscherba,  Tunis 

C.  racemosa  (Rudolphi) 

Coluber  sp.,  Ophiomorphus 

Brazil 

miliaris,  Ophis  merremii 

Brazil 

0.  trimeresurus  (Parona) 

Trimeresurus  formosus 

Island  of  Mentawei,   East 
Indies 

284 


ILLINOIS    BIOLOGICAL    MONOGRAPHS 


[284 


Families  of  Amphibia  and  Reptilia  Harboring  Proteocephalid  Parasites 


Order 
Family  Proteidae 

Necturtis  maculosus  Raf. 
Family  Amblystomidae 
Amhlystoma  tigrinum  (Green) 
Order 
Family  Hylidae 
Hyla  aurea 


Order 
Suborder 
Family  Varanidae 

Varanus  (Hydrosaurus)  salvator 
Varanus  varius  Shaw 
Varanus  gouldii  Gray 
Varanus  bellii 
Varanus  sp. 
Varanus  sp. 


AMPHIBIA 
Caudata  (Urodela) 

Ophiotaenia  lonnbergii  (Fuhrmann) 

Ophioiaenia  filar oides  La  Rue 
Salientia  (Anura) 

Ophiotaenia  hylae  Johnston 

REPTILIA 
Squamata  (Sauria) 
Sauria    (Lacertilia) 

Acanthotaenia  shipleyi  v.  Linstow 
Acanthotaenia  tidswelli  Johnston 
Acanthotaenia  tidswelli  Johnston 
Acanthotaenia  tidswelli  Johnston 
Acanthotaenia  biroi  (von  Ratz) 
Acanthotaenia  saccifera  (von  Ratz) 


Suborder      Serpentes   (Ophidia) 


Family  Boidae 
Subfamily  Boinae 
Eunectes  murinus  Wagl. 
Boa  constrictor 
Family  Colubridae 
Series  A.  Aglj'pha 
Subfamily  Colubrinae 
Natrix  rhombifer  Hallowell 
Natrix  sipedon 
Coluber  sp. 
Coluber  sp. 

Ophiomorphus  miliaris 
Ophis  merremii 
Series  B.  Opisthoglypha 
Subfamily  Homalapsinae 
Psammodynastes  pulverulentus 
Boie 
Series  C.     Proteroglypha 
Subfamily  Elapinae 
Pseudechis  porphyriacus  Shaw 
Pseudechis  australis  Gray 
Notechis  scutatus  Peters 
Denisonia  superba  Gtmther 


Crepidobothrium  gerrardii  (Baird) 
Crepidobothrium  gerrardii  (Baird) 


Ophiotaenia  perspicua  La  Rue 
Ophiotaenia  lactea  (Leidy)  sp.  inq. 
Ophiotaenia  nattereri  (Parona)        \/^ 
Ophiotaenia  racemosa  (Rudolphi) 
Ophiotaenia  racemosa  (Rudolphi) 
Ophiotaenia  racemosa  (Rudolphi) 


Ophiotaenia  pigmentata  (von  Linstow) 


Acanthotaenia  gallardi  Johnston 
Acanthotaenia  gallardi  Johnston 
Acanthotaenia  gallardi  Johnston 
Acanthotaenia  gallardi  Johnston 


285] 


PROTEOCEPHALIDAE—LA  RUE 


285 


REPTILIA 

Suborder      Serpentes  (Ophidia) 
Family  Viperidae 

Subfamily  Crotalinae 

Triineresurus  forinosus  Ophiotaenia   trimeresuri   (Parona) 

Lachesis  (Bothrops)  lanceolatush.   Ophiotaenia  calmettei  (Barrois) 
Ancistrodon  piscivorus  Holbr.  Ophiotaenia  marenselleri  (Barrois) 

Ancistrodon  piscivorus  Holbr.  Ophiotaenia  grandis  La  Rue 

Host   uijknown,    reported   to    have   come 
from  a  dog  but  probably  from  a  snake    Ophiotaenia  punica  (Cholodkovski) 


PROTEOCEPHALIDS  OF  FISH 

Hosts  and  Localities  of  Collection  of   Proteocephalid  Species  infesting  Fish  and 
the   Species   of   Pish   infested 


Cestode  Species 

Hosts 

Locality 

Proteocephalus    a- 

Alosa  finta  var.  lucustris 

Lake  Como,  Italy 

gonis 

P.  ambloplitis 

Ambloplites  rupestris 

Lake  George,  New  York 

Micro pterus  salmoides 

Lake  George,  New  York;  Walnut 
Lake,  Mich. ;  Pelican  Lake, 
Minn. 

Micropterus  dolomieu 

Lake  St.  Clair,  Mich.;  Lake  Men- 
dota,   Wis. 

Aiiiia  calva 

Lake  St.  Clair,  Mich. ;  Lake  Erie 

P.  cernuae 

Acerina  cernua 

Konigsberg,  Prussia;  Lake  Bala- 
ton, Hungary 

(?)A'cerina  schraetser 

Lake  Balaton,  Hungary 

P.  Cyclops  sp.  inq. 

Coregonus  maraena 

Schallsee,  Germany 

P.  dubius 

Perca  fluviatilis 

Lake  Geneva,  Switzerland 

P.  esocis 

Esox  lucius 

Reval,  Esthonia  (a  Baltic  prov- 
ince) 

P.  exiguus 

Coregonus  nigripinnis,   C. 

Lake   Michigan,   near  Charlevoix, 

prognathus,  C.  artedi 

Mich. 

P.  fallax 

Coregonus  maraena=(fera) 

Lake  Lucerne,  Switzerland 

P.  filicollis 

Gasterosteus  aculeatus 

Greifswald ;  Berlin ;  Paris ;  Ire- 
land 

G.  pungitius 

Rennes  ;  Halle ;  Greifswald ;  Re- 
val, Esthonia;  Porkhala,  Fin- 
land; Upsala,  Sweden. 

P.  fossatus 

Pimelodus  pati 

Rio  Paraguay,  S.  A. 

P.  hemisphericus 

Anguilla  vulgaris 

Padua,  Italy 

sp.  inq. 

P.  longicollis 

Coregonus  wartmanni    / 

? 

C.  wartmanni  nobilis 

Lake    Lucerne 

286 


ILLINOIS    BIOLOGICAL    MONOGRAPHS 


[286 


Cestode  Species 

Hosts 

Locality 

P.  longicollis 

C.    maraena=(C.    lavaretus 

Greifswald;  Rositten,  E.  Prussia; 

and  C.  fera) 

Lake  Geneva 

C.  maraenula 

Berlin 

C.  albula 

Nikolaiken,  E.  Prussia 

C.  schinzii  helveticus 

Lake  Lucerne 

C.  exiguus  albellus 

Lake  Lucerne 

• 

Osmervs  eperlanus 

Berlin;  locality  not  recorded  sev- 
eral times 

Trutta  trutta 

Germany    ( ?) 

T.  fario 

? 

Saint 0   thymallus 

? 

S.  salveHnus=S.  alpinus  and 

Lake  Lucerne ;  Lake  Geneva ;  lo- 

S. umbla 

cality  not  stated 

Very  doubtful  hosts!!!! 

(?)  Esox  Indus 

Lake  Lucerne;  Lake  Balaton, 
Hungary 

{?)Perca  fluviatilis 

Lake  Lucerne 

(?)  Albumus  lucidus 

Lake  Lucerne 

_. 

(?)  Squalius  cephalus 

Lake  Lucerne 

(?)Leuciscus  leuciscus 

Lake  Lucerne 

P.  macrocephalus 

Anguilla  vulgaris 

Greifswald ;  Memel,  East  Prussia ; 
Rennes ;  Padua ;  Narenta ; 
Trieste;  Lake  Lucerne;  Finland 

Anguilla  chrysypa 

Wood's  Hole,  Mass. ;  Sebago 
Lake,  Maine 

P.  macrophallus 

Cichla  monoculus  Spix 

Brazil 

sp.  inq. 

P.  neglectus 

"Forelle",  Trutta  fario 

Lake  Lucerne  and  Lake  Geneva, 
probably 

P.  nematosoma  sp. 

Esox  reticulatus 

Locality     not     known,     probably 

inq. 

eastern  United  States 

P.  osculatus  sp. 

Silurus  giants 

Magdeburg  and  Greifswald 

inq. 

P.  pentastoma 

Polypterus  bichir 

Khor  Attar  on  White  Nile 

P.  percae 

Perca  norvegica 

? 

Perca  fluviatilis 

Greifswald;  Rositten,  East  Prus- 
sia ;  Finland ;  Upsala 

Coregonus  viaraena={C. 

Finland 

lavaretus) 

{})Acerina  cernua 

? 

(  ?)Cottus  quadricomis 

Finland 

(?)Gasterosteus  aculeatus 

Rositten,  East  Prussia 

P.  perplexus 

Aitiia  calva  and  Lepisosteus 
platostomus 

Illinois  River,  Havana,  Illinois 

287] 


PROTEOCEPHAUDAE—LA  RUE 


287 


Cestode  Species 

Hosts 

Locality 

P.  pinguis 

Esox  reticulatus 

Sebago  Lake,  Maine 

Esox  lucius 

Walnut  Lake,  Michigan 
Lake  Geneva,  Wisconsin 

P.  pusillus 

Saint  0  sebago 

Sebago  Lake,  Maine 

Cristivomer  namaycush 

Lake  Temagami,  Ontario 

Proteocephalus  sa- 

Cobitis  barbatula 

Petersburg  and  Province  of  Nov- 

gittus sp.  inq. 

gorod,  Russia 

P.  salmonis-umblae 

Salmo  salveUnus=^{S.  um- 

Lake    Geneva,    Switzerland,    Ger- 

sp. inq. 

bla) 

many 

P.  salvelini  sp.  inq. 

Cristivomer  namaycush 

Outer  Island,  Lake  Superior 

P.  singularis 

Lepisosteus  platostomus 

Illinois  River,  Havana,  111. 

P.  skorikowi 

Acipenser  stellatus 

River  Gurgen,  Caspian  Sea 

P.  sulcatus 

Polypterus  endlicheri 

Duem  on  the  White  Nile 

Clarotes  laticeps 

Khartoum  on  the  White  Nile 

P.  torulosus 

Idus    melanotus={Cyprinus 
jeses  and  Leuciscus  idus) 

Berlin;  Greifswald;  Finland 

Leuciscus  leuciscus= 

Germany     ( ?)  ;     Podiebrad,     Bo- 

(Squalius leuciscus) 

hemia  ;    Lake    Lucerne 

L.  grislagine 

Locality    (?) 

Alburnus  bipnnctatus 

?  (Material  in  M.  C.  V.) 

A.   sp. 

Germany    (?)    (from    context   of 
report) 

A.  lucidus 

Lakes  Geneva  and  Lucerne 
Germany    (?)    (from   context   of 
report) 

Aspius  rapax 

Germany    (?)     (from    context   of 
report) 

Abramis  brama 

Lake  Balaton,  Hungary 

Pelecus  cultratus 

Lake  Balaton,  Hungary 

Gobio  fluviatilis 

Lake  Lucerne 

Blicca  bjoerkna 

Lake  Lucerne 

Very  doubtful  hosts  ! ! ! ! ! 

{DCoregonus  maraena 

Lake  Lucerne 

(fera) 

(?)C  exiguus  albellus 

Lake  Lucerne 

(?)C.  schinsii  helveticus 

Lake  Lucerne 

(?)  Salmo  salvelinus 

Lake  Lucerne 

(?)Perca  fluviatilis 

Lake  Lucerne  and  Podiebrad,  Bo- 
hemia 

(?)Lota  vulgaris 

Lake  Geneva 

Corallobothriuvt 

Malopterurus  electricus 

Egypt 

solidum 

C.  lobosum 

Silurus  sp. 

Rio  Paraguay,  South  America 

Choanoscolex  ab- 

Silurus  sp. 

Rio  Paraguay,  South  America 

scisa 

288  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [288 

Distribution  of  the  Proteocephalids  of  Fish  According  to 

Continents 

The  accompanying  table  lists  the  Proteocephalid  species  which  are 
found  on  each  continent.  An  examination  of  the  table  shows  that  in 
Europe  there  are  12  well  known  and  5  doubtful  species.  All  are  pecu- 
liar to  Europe  with  the  exception  of  P.  macrocephalus  which  occurs 
also  in  North  America  along  the  Atlantic  coast.  In  North  America  9 
species  are  known.  Of  these  2  are  doubtful.  All  are  peculiar  except 
P.  macrocephalus.  From  South  America  4  species  are  recorded.  All 
are  peculiar.  Likewise  from  Africa  3  species  are  recorded  and  all  are 
peculiar.  No  species  of  this  group  have  yet  been  reported  from  Asia 
or  Australia,  probably  because  but  little  parasitological  work  has  been 
done  on  the  fish  of  those  continents.  Nor  are  there  any  records  as 
yet  from  the  West  or  East  Indies,  Greenland,  or  Iceland.  There 
has  been  but  a  singe  collection  record  from  Great  Britain,  where  the 
parasites  of  fresh- water  fishes  seem  to  have  had  little  attention. 

The  records  are  as  yet  too  meager  to  justify  any  broad  conclusions 
as  to  the  distribution  of  these  cestodes  over  the  larger  land  areas.  One 
may  expect  to  find  the  same  Proteocephalid  species  in  the  hosts  which 
are  common  to  Europe  and  North  America.  One  such  case  is  known, 
namely,  that  of  P.  macrocephalus  in  Anguilla.  Esox  lucius  which  is 
common  to  both  of  these  continents  harbors  one  species  in  Europe  and 
another  one  in  North  America.  Other  fish  species  occurring  in  both 
continents  have  apparently  yielded  no  records  in  the  literature. 

Distribution  of  Fish  Proteocephalids  according  to  Continents 


Continent 
Europe 


North  America 


South    America 

Africa 

Asia 

Australia 


Species  of  Proteocephalid  found 


Proteocephalus  agonis,  P.  cernuae,  P.  cyclops  sp.  inq.,  P.  dubitis, 

P.  esocis,  P.  fallax,  P.  filicollis,  P.  hemisphericus  sp.  inq.,  P. 

longicoUis,  P.  macrocephalus,  P.  neglectus,  P.  osculatus  sp.  inq., 

P.  percae,  P.  sagittus  sp.  inq.,  P.  salmonis-umblae  sp.  inq.,  P. 

skorikowi,  P.  torulosus. 
P.  ambloplitis,  P.  exiguus,  P.  macrocephalus,  P.  nematosoina  sp. 

inq.,  P.  perplexus,  P.  pinguis,  P.  pusillus,  P.  salvelini  sp    inq., 

P.  singularis. 
P.  fossatus,  P.  macrophallus  sp.  inq.,  Corallobothrium  lobosum, 

Choanoscolex  abscisa. 
P.  pentastoma,  P.  sulcatus,  Corallobothrium  solidum. 
No  species  reported. 
No  species  reported. 


289]  PROTEOCEPHALIDAE—LA  RUE  289 

Fish  Haeboring  Two  or  More  Proteocephalid  Species 

An  examination  of  the  complete  list  of  fish  Proteocephalids  shows 
that  there  are  fourteen  host  species  which  are  reported  to  harbor  more 
than  one  species  of  Proteocephalus.  The  data  relating  to  these  cases 
are  presented  in  the  accompanying  table  which  names  the  hosts,  their 
parasites  and  the  distribution  of  the  latter.  A  brief  study  of  the  table 
shows  that  of  the  fourteen  host  species  ten  are  accredited  with  two 
species  of  Proteocephalus  each,  three  with  three,  and  one  host  with  four. 
If  from  these  numbers  are  deducted  the  instances  of  doubtfully  identi- 
fied species  and  the  species  inquirenda  there  follow  these  results:  Five 
host  species  each  harbor  two  well  described  Proteocephalus  species  while 
one  host  harbors  three.  There  are  then  but  six  undoubted  cases  of  two 
or  more  species  of  Proteocephalus  occurring  in  the  same  host  species. 

An  analysis  of  these  six  cases  shows  that  in  Lepisosteus  platostomus 
the  two  parasitic  species  occur  in  the  same  locality,  while  in  the  cases 
of  Amia  calva,  Esox  lucius,  and  Perca  fluviatilis  the  two  parasitic  species 
each  occur  in  different  localities.  Likewise  the  three  species  of  Proteo- 
cephalus found  in  Coregonus  maraena  occur  each  in  different  localities. 
The  distribution  of  the  parasites  of  Trutta  fario  is  not  known.  One 
may  conclude  that  two  or  more  species  of  Proteocephalus  may  inhabit 
the  same  host  species  in  the  same  locality  or  in  different  localities.  The 
factors  which  determine  this  are  not  here  shown. 


Species  of  Fish  which  harbor  More  than  One  species  of  Proteocephalus 


Host 

Proteocephalus    species 
harbored 

Distribution  of  latter 

Amia  calva 

Proteocephalus  perplcxus 

Illinois  River 

P.  ambloplitis 

Lake  St.  Clair ;  Lake  Erie 

Lepisosteus  plato- 

P. singularis 

Illinois    River 

stomus 

P.  perplexus 

Illinois   River 

Salmo  salvelinus 

P.  longicollis 

Lakes   Geneva  and   Lucerne,   and 
locality  not  known 

P.  salmonis-ufitblae  sp.  inq. 

Lake  Geneva ;  Germany 

P.  torulosus  (?) 

Lake  Lucerne 

Trutta  fario 

P.  longicollis 

Locality  not  known 

P.  neglectus 

Lake  Geneva  and  Lake  Lucerne 

Cristivomer  namay- 

P.  pusillus 

Lake  Temagami,  Ontario 

cush 

P.  salvelini  sp.  inq. 

Lake   Superior 

290 


ILLINOIS    BIOLOGICAL    MONOGRAPHS 


[290 


Species  of  Fish  which  harbor  More  than  One  species  of  Proteocephalus  (Continued) 

Host 

Proteocephalus    species 
harbored 

Distribution  of  latter 

Coregonus  maraena 

P.  longicollis 

Greifswald;   Rositten,  East  Prus- 

(C.   lavaretus     and 

sia;  Lake  Geneva 

C.  fera) 

P.  fallax 

Lake  Lucerne 

P.  percae 

Finland 

P.  torulosus  (?) 

Lake  Lucerne 

Alburnus  lucidus 

P.  torulosus 

Germany  (  ?)  ;  Lake  Lucerne  and 
Lake   Geneva 

P.  longicollis  (  ?) 

Lake  Lucerne 

Leuciscus  leuciscus 

P.   torulosus 

Germany  (?);  Bohemia;  Lake 
Lucerne 

P.  longicollis  (?) 

Lake  Lucerne 

Anguilla  vulgaris 

P.  macrocephalus 

Finland ;  Germany ;  France ; 
Switzerland ;  Hungary ;  Italy 
(Padua) 

P.  hemisphericus  sp.  inq. 

Padua,  Italy 

Esox  lucius 

P.  esocis 

Finland 

P.  pinguis 

Maine;  Michigan;  Wisconsin 

P.  longicollis  (?) 

Lake  Lucerne;  Lake  Balaton, 
Hungary 

Esox  reticulatus 

P.  pinguis 

Sebago  Lake,  Maine 

P.  nematosoma  sp.  inq. 

Eastern     (?)    United    States 

Gasterosteus  aculeatus 

P.  filicollis 

Greifswald  and  Berlin,  Germany; 
Paris;   Ireland 

P.  percae   (?) 

East  Prussia 

Perca  fluviatUis 

P.  percae 

Germany;  Finland;   Sweden 

P.   dubius 

Lake  Geneva 

P.  longicollis   (?) 

Lake   Lucerne 

Acerina  cernua 

P.  cernuae 

Prussia ;  Lake  Balaton,  Hungary 

P  percae  (?) 

Locality  unknown 

Proteocephalids  Occurring  in  Two  or  More  Species  op  Fish 


Twelve  Proteocephalid  species  have  been  found  to  occur  in  more 
than  one  host  of  more  or  less  close  relationship.  These  species  are  as 
follows :  Proteocephalus  amhloplitis,  P.  cernuae,  P.  exiguus,  P.  filicollis, 
P.  longicollis,  P.  macrocephalus,  P.  percae,  P.  perplexus,  P.  pinguis, 
P.  pusillus,  P.  sulcatus,  P.  torulosus.  In  the  following  pages  each  of 
these  species  is  discussed  separately.  The  hosts  have  been  investigated 
as  to  distribution  and  as  to  food  habits  in  the  hope  that  some  clue  might 
be  found  to  the  relations  existing  between  host  and  parasite,  a  relation 


291]  PROTEOCEPHALIDAE—LA  RUE  291 

which  in  some  cases  is  far  different  from  that  expected  if  the  idea  of 
the  specificity  of  hosts  be  strictly  adhered  to.  The  data  on  distribution 
and  food  habits  of  the  hosts  have  been  abstracted  and  combined  from 
several  sources,  chief  of  which  are  Nitsche  (1909),  Leunis  (1883), 
Bridge  (1904)  and  Boulenger  (1904)  in  Cambridge  Natural  History, 
volume  VII,  Jordan  and  Evermann  (1896-1900),  and  Forbes  and  Rich- 
ardson (1909). 

Proteocephalus  amhlopUtis 
This  species  is  known  at  present  only  from  waters  which  drain  into 
the  St.  Lawrence  and  the  Red  River  of  the  North.  Altho  Professor 
Ward  examined  the  proper  host  species  from  the  Illinois  River  at 
Havana,  111.,  during  the  summer  of  1909  his  collections  show  that  this 
parasite  was  not  present  there.  The  reported  hosts  are  AmhlopUtis 
rupestris,  Micropterus  salmoides,  M.  dolomieu,  and  Amia  calva.  The 
first  three  hosts  have  much  the  same  distribution  over  eastern  and  cen- 
tral North  America  to  the  Gulf  of  Mexico,  while  Amia  calva  occurs 
over  a  somewhat  more  restricted  area.  It  is  abundant,  however,  over  a 
large  part  of  the  area  in  which  the  other  hosts  are  found.  In  many 
localities  two,  three,  or  even  four  of  the  hosts  live  in  the  same  waters. 
All  four  hosts  are  carnivorous,  eating  other  fish  and  crawfish.  Some  of 
them  eat  other  food  in  addition.  These  four  hosts  live  in  the  same 
general  region,  frequently  in  the  same  waters,  and  have  certain  food 
habits  in  common.  Such  are  the  conditions  to  be  expected  if  the  hosts 
are  to  harbor  a  common  parasitic  species.  One  question  remaining  to 
be  solved  is  this:  why  is  this  species  not  found  in  the  Illinois  River 
where  these  hosts  live  in  large  numbers?  Since  the  Illinois  River  was 
connected  with  the  Great  Lakes  in  the  recent  geological  period  the 
failure  to  find  this  species  in  this  river  seems  the  more  remarkable. 

Proteocephalus  cernuae 

P.  cernuae  is  known  at  present  only  from  Prussia  and  from  Lake 
Balaton,  Hungary.  The  waters  of  Prussia  drain  to  the  Baltic  Sea  while 
those  of  Lake  Balaton  drain  into  the  Danube  and  so  to  the  Black  Sea. 
The  hosts  of  this  species  are  Acerina  cernua  and  A.  schraetzer.  The 
former  has  a  wide  distribution  in  north  and  central  Europe  and  is  also 
found  in  the  Danube.  A.  schraetzer  is  confined  to  the  Danube  and  its 
tributaries.  Thus  in  the  Danube  the  ranges  of  the  two  species  overlap. 
Both  species  are  carnivorous  and  both  take  bottom  fauna.  Under  these 
conditions  of  distribution  and  food  habits  one  would  expect  to  find  both 
species  harboring  the  same  cestode  species.  With  further  study  of  the 
collections  this  parasite  will  probably  be  found  to  have  a  much  wider 
and  a  continuous  distribution. 


292  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [292 

Proteocephalus  exiguus 
P.  exiguus  has  been  found  only  in  the  Great  Lakes.  Its  hosts, 
Coregonus  (Argyrosomus)  artedi,  C.  nigripinnis,  and  C.  prognathus  are 
all  found  in  certain  of  the  Great  Lakes.  C.  artedi  is  in  all  the  Great 
Lakes  and  neighboring  waters,  and  north  to  the  Arctic  regions,  exclu- 
sive of  Alaska.  C.  nigripinnis  is  found  in  the  small  lakes  of  Wisconsin 
and  Minnesota  and  in  Lake  Michigan.  C.  prognathus  occurs  probably 
in  all  the  Great  Lakes  in  deep  water.  Thus  in  Lake  Michigan,  at  least, 
these  species  are  all  found.  As  to  food  habits  but  few  data  may  be 
found  for  C.  prognathus  and  C.  7iigripinnis.  C.  artedi  feeds  on  Ento- 
\/'  mostraca,  small  molluscs  and  crustaceans,  with  larvae  of  insects  and 
other  animal  forms  including  bottom  fauna  and  spawn  of  the  common 
white  fish.  It  seems  probable  that  the  other  species  take  many  of  these 
same  food  elements.  Under  these  conditions  the  finding  of  this  species 
in  three  closely  related  hosts  is  not  remarkable  but  is  to  be  expected. 

Proteocephalus  filicollis 
This  species  is  known  only  from  the  lower  reaches  of  the  streams 
flowing  into  the  Baltic  Sea,  into  the  North  Sea  and  the  waters  adjacent 
thereto.  It  apparently  does  not  occur  in  the  Alpine  lakes.  Its  hosts 
are  Gasterosteus  aculeatus  and  G.  pungitius.  The  latter  is  found  all 
over  north  Europe  (not  in  south  Germany)  and  in  North  America  as 
far  southward  as  the  Great  Lakes  region.  G.  aculeatus  has  a  more 
restricted  distribution  being  confined  to  the  fresh  and  brackish  or  saline 
waters  of  Europe  with  the  exception  of  the  Danube  drainage  system. 
Thus  the  two  species  occupy*  north  Europe  in  common.  In  food  habits 
the  species  are  much  alike.  Both  are  voracious  plankton  feeders;  both 
take  also  fish-eggs  and  fry,  and  bottom  fauna.  These  are  the  conditions 
which  one  would  expect  to  find  if  the  two  species  harbor  the  same  para- 
site species.  Specimens  of  G.  pungitius  from  North  America  should  be 
examined  in  order  to  learn  whether  they  harbor  the  same  species  of 
parasite  as  the  European  specimens. 

Proteocephalus  longicollis 

This  species  has  a  wide  distribution  in  Germany  and  is  known  also 
from  Switzerland.  It  is  unknown  in  southern  Europe,  Russia  or  in 
France.  It  has  been  reported  from  Hungary,  probably  as  the  result  of 
\y  a  misdetermination.  Its  reported  hosts  are  mainly  salmonids  tho  the 
V  list  includes  one  esocid,  one  percid,  and  three  cyprinidg.  The  accom- 
panying table  names  the  hosts  and  shows  the  distribution  and  food  of 
each.    An  examination  of  the  table  shows  that  in  a  general  way  these 


293]  PROTEOCEPHALIDAE—LA  RUE  293 

host  species  overlap  greatly  in  their  distribution.  Several  of  the  species 
occur  in  the  same  waters.  Some  live  only  in  the  depths  of  deep  lakes 
while  others  live  only  on  the  surface ;  others  live  in  brooks  and  streams, 
still  others  as  Perca  fiuviatilis  are  ubiquitous.  One  host,  Trutta  trutta, 
lives  in  salt  water  and  enters  fresh  water  to  spawn.  Thus  tho  a  species 
may  have  a  wide  distribution  its  habitat  may  be  narrowly  restricted. 
In  the  restricted  habitat  the  food  of  the  species  is  likely  to  be  more  or 
less  restricted.  Surface  fish  do  not  get  the  same  kinds  of  food  as  do 
those  that  live  on  the  bottom. 

A  survey  of  the  table  shows  that  all  the  hosts  of  P.  longicollis  de- 
pend on  animal  food  tho  some  of  them  may  take  plant  food  on  occasion. 
Certain  species  are  much  restricted  as  to  food,  so  much  restricted,  in- 
deed, that  it  may  be  s^id  with  some  degree  of  certainty  that  some  of  the 
plankton  forms  must  be  carriers  of  this  infection.  Coregonus  wart- 
manni  and  C.  wartmanni  nohilis  are  referred  to  as  hosts  which  eat 
plankton  exclusively.  It  is  also  apparent  that  if  the  records  be  true 
some  other  invertebrates  must  be  carriers  of  the  infection  for  several 
species  are  found  which  do  not  feed  on  plankton.  Some  molluscs  may 
play  a  part  as  carriers.  Still  other  fish  of  the  list  eat  fish  almost  ex- 
clusively hence  it  must  be  that  fish  can  serve  as  intermediate  hosts 
for  this  parasite.  It  has  already  been  shown  in  numerous  instances 
that  the  primary  host  may  also  serve  as  the  intermediate  host  for 
Proteocephalid  species.  If  one  considers  now  the  different  habitats  and 
the  various  kinds  of  food  taken  in  them  it  must  be  admitted  that  the 
evidence  points  toward  a  confusion  of  several  species  under  the  one 
name.  In  the  opinion  of  the  writer  the  validity  of  many  of  the  records 
is  doubtful.  A  comparative  study  of  Proteocephalids  taken  from  all 
these  hosts  of  different  localities  would  perhaps  settle  these  questions 
as  to  hosts  of  P.  longicollis. 

Granting  that  all  the  salmonids  of  the  list  are  hosts  of  this  parasite 
then  on  the  basis  of  food,  distribution,  and  habitats  it  seems  that  the 
non-salmonids  might  likewise  be  its  hosts.  However,  it  has  seemed 
that  these  records  relating  to  the  non-salmonids  rest  on  too  slender  a 
foundation.  Without  an  exception  these  records  are  those  of  Nufer 
(1905)  whose  work  has  been  discussed  in  the  section  dealing  with  the 
description  of  P.  longicollis.  Parasites  of  these  non-salmonids,  espe- 
cially from  Lake  Lucerne  where  Nufer  was  at  work,  ought  to  be  sub- 
jected to  a  careful  comparative  study.  This  would  doubtless  establish 
or  overthrow  the  validity  of  the  records. 


294 


ILLINOIS    BIOLOGICAL    MONOGRAPHS 


[294 


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PROTEOCEPHALIDAE—LA  RUE 


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296  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [296 

Proteocephalus  macrocephalus 
This  species  has  the  widest  distribution  of  any  known  Proteocepha- 
lid.  It  has  been  taken  in  many  localities  of  northern,  central  and  south- 
ern Europe  and  from  two  localities  in  eastern  North  America.  Its  hosts 
are  Anguilla  vulgaris  and  A.  chrysypa.  The  former  has  a  wide  range 
over  most  of  Europe,  north  Africa  and  temperate  Asia.  It  is  lacking 
in  the  Black  and  Caspian  Seas.  By  some  authorities  this  species  is 
thought  to  be  found  also  in  North  America.  Others,  including  Jordan 
and  Evermann  (1896-1900)  make  the  American  form  a  separate  species, 
calling  it  A.  chrysypa.  This  species  occurs  over  the  most  of  North 
America  east  of  the  Rocky  Mountains,  including  Mexico  and  the  West 
Indies.  The  food  habits  of  the  two  species  are  alike.  They  are  vora- 
cious, and  act  as  scavengers,  preferring  dead  fish  or  other  animal  refuse. 
In  view  of  their  close  relations  and  their  similar  habits  it  is  not  surpris- 
ing that  these  species  harbor  the  same  species  of  parasite. 

Proteocephalus  percae 
The  geographical  distribution  of  this  species  is  imperfectly  known. 
Up  to  the  present  time  it  has  been  taken  only  in  East  Prussia,  Finland 
'and  Sweden.  It  has  not  yet  been  reported  from  the  Swiss  lakes  tho 
two  of  its  host  species  are  abundant  in  that  region.  These  hosts  are 
Perca  fluviatUis  and  Coregonus  maraena  (fera).  The  appended  table 
gives  the  data  as  to  hosts,  their  distribution  and  food  habits.  An  exam- 
ination of  the  table  shows  certain  points  of  interest.  Perca  marina 
which  Miiller  gave  as  a  host  of  this  parasite  is  little  known,  nor  is  Perca 
norvegica  which  Rudolphi  said  was  the  host  of  Miiller 's  parasite.  The 
remaining  host  species  overlap  in  their  distribution.  Likewise  their 
food  habits  overlap  to  a  certain  extent.  All  eat  bottom  fauna  tho  Perca 
ftuviatUis  preys  more  on  other  fishes  than  do  the  other  hosts.  The  food 
habits  of  P.  fluviatUis  and  Coregonus  maraena  show  close  resemblance. 
These  two  species  are  known  to  harbor  the  species  which  Schneider 
(1905)  describes  as  Ichthyotaenia  percae  (Miiller).  Acerina  cernua 
and  Qasterosteus  aculeatus  may  harbor  this  parasite  but  it  seems  doubt- 
ful. These  two  species  are  known  to  harbor  Proteocephalus  cernuae 
and  P.  filicollis  respectively.  It  seems  that  there  may  have  been  a  mis- 
determination.  The  parasites  of  these  two  hosts  and  of  Coitus  quadri- 
carnis  should  be  subjected  to  a  re-examination. 


297] 


PROTEOCEPHALIDAE—LA  RUE 


2ff7 


Hosts  of  Proteocephalus  percae,  their  Food  and  Distribution 

Host  species 

Distribution 

Food    habits 

Perca  norvegica=Perca  ma- 

No data 

Xo  data 

nna 
Perca  fluviatilis 

Lakes  and  rivers  of  all  Eur- 

Carnivorous,      eats       fish 

ope  and  northern  Asia 

(largely    species    of    Al- 
bumus),  amphibia,  snails, 
insects   and  worms 

Coregonus    maraena=(.C. 

In  deep  water  of  large  lakes 

Small  "fish,    bottom    fauna. 

lavaretus  and  C.  fera) 

of  Alps,  Bavaria,  and  near 

Plankton,  small  Crustacea 

Baltic  coast;  Baltic  Sea 

and  molluscs 

Acerina  cemua 

In  brackish  and  fresh  wa- 

Fish eggs  and   fry,  bottom 

ters  of  north  and  central 

fauna,  rooting  in  the  bot- 

Europe    to     Siberia;     in 

tom  for  last 

ENanube.       Prefers     deep 

water  with  sandy  bottom 

Gasterosteus  aculeatus 

In    fresh    and    brackish    or 

A     plankton     feeder;     eats 

salt  water  of  all  Europe 

fish-eggs     and     fry,     also 

except  Danube 

bottom    faima 

Coitus  quadricomis 

Arctic  regions  south  to  Bal- 

No data  on  food.    A  close 

tic      Sea,     westward      to 

relative,     C.    gohio,    eats 

Greenland 

molluscs  and  spawn. 

Proteocephalus  perplexus 

This  species  is  known  at  present  only  from  the  Illinois  River  at 
Havana,  111.  Leidy's  notice  (1886)  of  Taenia  fiZicoUis  found  in  Amia 
calva  from  North  Carolina  may  refer  to  this  species.  The  hosts  are 
Amia  calva  and  Lepisosteus  platostomus.  The  former  is  common  in 
rivers  and  lakes  of  central  and  southern  North  America,  including  lakes 
Huron  and  Erie.  The  latter  host,  Lepisosteus  platostomus,  occurs 
throughout  the  Mississippi  valley  but  is  said  to  avoid  the  smaller 
streams.  Thus  in  the  Mississippi  valley  the  two  hosts  have  a  common 
range.  Amia  feeds  on  fish,  crawfish  and  molluscs  while  the  other  host 
takes  fish  almost  exclusively.  If  Lepisosteus  includes  Amia  in  its  diet 
it  may  secure  its  infection  from  Amia  which  may  perhaps  serve  as  both 
primary  and  secondary  host.  On  account  of  its  armor  it  is  not  at  all 
likely  that  the  gar  serves  as  food  for  Amia.  It  has  seemed  rather  re- 
markable that  this  parasite  was  not  found  in  Amia  calva  of  the  Great 
Lakes.  A  most  careful  examination  of  the  material  collected  by  Pro- 
fessor Ward  in  the  course  of  his  work  on  the  Great  Lakes  has  shown 
that  it  does  not  exist  there.  Likewise  it  seems  remarkable  that  the  other 
carnivorous  fish  of  the  Illinois  River,  such  as  the  bass,  do  not  have  this 
infection.    Some  of  these  fish  eat  very  much  the  same  food  as  Amia. 


298  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [298 

Proteocephalus  pinguis 

P.  pinguis  is  a  North  American  species  found  in  widely  separated 
localities.  It  has  been  found  in  small  lakes  of  Maine,  Michigan  and 
Wisconsin,  It  has  not  yet  been  found  in  the  Great  Lakes.  Further 
collections  may  show  a  much  wider  distribution  for  this  species.  The 
hosts,  Esox  lucius  and  E.  reticulatus,  have  a  wide  distribution.  Esox 
lucius  is  a  cosmopolitan  species  of  the  northern  hemisphere,  occurring 
in  the  fresh  waters  of  Europe,  Asia  and  North  America.  In  North 
America  its  range  is  from  Alaska  south  to  the  upper  Mississippi  valley, 
east  of  the  Rockies  to  the  Potomac.  Esox  reticulatus  is  common  east 
and  south  of  the  Alleghanies  to  Louisiana,  and  Arkansas.  The  ranges 
of  these  hosts  are  common  east  of  the  Alleghanies  to  the  Potomac  river. 

Both  hosts  are  voracious  carnivores,  eating  fish,  frogs,  crawfish, 
mice,  reptiles,  water  birds,  and  larger  insects.  That  these  two  host  spe- 
cies harbor  the  same  species  of  Proteocephalus  is  then  not  remarkable. 
Since,  however,  Esox  lucius  is  common  to  Europe  and  North  America 
one  might  expect  to  find  it  harboring  the  same  species  of  Protecephalus 
in  both  continents.  Such  does  not  seem  to  be  the  case.  This  fact  sug- 
gests that  these  two  species,  P.  pinguis  from  North  America  and  P. 
esocis  found  in  the  European  Esox,  have  arisen  as  varieties  of  the  same 
species  or  that  the  host  upon  going  into  the  new  region,  either  Europe 
or  North  America,  has  acquired  a  new  species  of  parasite. 

Proteocephalus  pusUlus 

This  species  occurs  in  Sebago  Lake,  Maine,  and  Lake  Temagami 
Ontario.  Its  hosts  are  Salmo  sehago  and  Cristivomer  namaycush. 
These  two  salmonids  live  ou^y  in  fresh  water  and  their  habitats  overlap 
to  a  certain  extent.  Salmo  sehago,  according  to  Jordan  and  Evermann 
(1896),  occurs  in  Sebago  Lake  and  northward,  in  lakes,  rarely  entering 
streams.  Cristivomer  namaycush  lives  in  the  lakes  of  New  York,  New 
Hampshire  and  Maine  and  westward  to  the  headwaters  of  the  Columbia 
and  north  to  the  Arctic  circle  (Forbes  and  Richardson,  1909).  Both 
fish  are  predaceous,  living  largely  on  fish.  Professor  "Ward's  records 
show  that  in  Sebago  Lake  Salmo  sehago  was  feeding  on  Osmerus  mor- 
dax.  Records  are  too  meager  to  allow  the  making  of  any  general 
conclusions. 

Proteocephalus  sulcatus 

This  species  is  known  only  from  the  Nile  River,  from  two  widely 
separated  hosts,  namely  Polypterus  endlicheri  and  Clarotes  laticeps. 
The  former  is  one  of  the  Polypteridae,  a  Crossopterygian,  while  the 
latter  named  host  is  a  Silurid,  of  the  order  Teleostei.  Both  hosts  occur 
in  the  Nile.  The  food  habits  of  Polypterus  endlicheri  are  not  well 
known.    One  species  of  Polypterus  feeds  on  small  teleosts  while  others 


299]  PROTEOCEPHALIDAE—LA  RUE  299 

take  batrachians  and  crustaceans  in  addition  to  the  fish.  This  species 
is  no  doubt  carnivorous  in  habits.  The  writer  is  unable  to  find  anything 
about  the  food  of  Clarotes  laticeps.  It  seems  probable,  however,  that 
it  is  a  carnivorous  species  as  are  many  others  of  the  Siluridae.  That 
this  parasite  is  able  to  find  the  proper  conditions  of  life  in  two  species 
as  widely  separated  as  are  these  seems  remarkable  altho  not  more 
remarkable  than  that  Amia  calva  may  serve  as  the  host  of  a  species 
characteristic  of  the  Centrarchidae.  It  has,  however,  been  pointed  out 
in  the  descriptive  part  of  the  work  on  this  species  that  there  seems  to 
be  some  reason  for  believing  that  two  species  have  been  confused  under 
the  one  name. 

Proteocephalus  torulosus 

This  species  is  known  only  from  northern  and  central  Europe, 
chiefly  from  the  Baltic  drainage.  Its  hosts  are  chiefly  cyprinids.  All  U^ 
the  earlier  reports  record  it  only  from  cyprinids.  During  the  last  few 
years  it  has  been  reported  from  Salmonids,  four  species,  and  from  Perca  '"" 
fluviatilis  and  Lota  vulgaris.  A  table  is  appended  showing  the  names 
of  the  hosts  as  reported,  their  distribution,  and  their  food.  As  to  food 
the  data  are  very  fragmentary  and  not  specific  yet  they  are  of  interest. 
An  examination  of  the  table  shows  that  the  hosts  are  common  in  the 
waters  of  the  lakes  and  streams  of  central  and  northern  Europe.  Some 
hosts  are  of  wide  distribution,  others  more  or  less  restricted.  Many  of 
these  hosts  occupy  the  same  waters,  some  as  surface  dwellers,  others  as 
bottom  dwellers,  some  in  the  deep  lakes,  others  in  the  small  streams,  while 
still  others  occur  in  the  brackish  bays  and  inlets  of  the  Baltic  coast. 
Thus  the  habitats  are  quite  varied. 

The  food  taken  by  the  host  varies  to  a  large  degree  with  the  habitat 
which  the  host  prefers.  Thus  from  the  table  it  is  found  that  the  food 
habits  of  the  hosts  listed  are  very  different.  Some  are  exclusively 
plankton  feeders,  others  feed  largely  on  bottom  fauna  or  bottom  fauna 
with  plankton.  Still  other  hosts  are  largely  eaters  of  fish.  If  the  food 
of  the  two  groups  of  hosts,  cyprinids  and  non-cyprinids,  be  considered 
it  is  noted  that  each  of  the  latter  group  takes  several  kinds  of  food. 
This  statement  applies  only  to  those  non-cyprinids  the  food  of  which 
is  known.  Each  of  the  cyprinids,  however,  takes  but  few  kinds  of  food. 
The  table  speaks  for  itself.  The  cyprinids  of  the  table  are  chiefly  feed- 
ers on  bottom  fauna.  Plankton  forms  a  large  part  of  the  food  of  three, 
and  fish  of  two  of  these  hosts.  Certain  elements  in  the  food  such  as  air- 
borne insects,  birds,  plants  and  probably  also  amphibians  (semi-terres- 
trial forms  especially)  do  not  function  as  intermediate  hosts  for  Pro- 
teocephalus species.  For  this  reason  they  need  not  enter  into  this 
discussion. 


JOO 


ILLINOIS    BIOLOGICAL    MONOGRAPHS 


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302  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [302 

How  can  the  infection  of  so  many  species  of  fish  with  this  parasite 
be  explained?  For  the  sake  of  the  argument  it  is  granted  that  all 
these  species  are  suitable  hosts  for  this  parasite.  If  all  these  species  are 
to  be  infected  it  is  necessary  that  some  of  the  invertebrates  included 
among  plankton  forms,  bottom  fauna,  and  molluscs  must  be  able  to  serve 
as  intermediate  hosts  of  this  parasite.  The  fish  which  become  infected 
from  eating  these  invertebrate  hosts  may  likewise  become  self-infected 
and  so  serve  as  intermediate  hosts.  If  such  fish  are  then  eaten  by  the 
predaceous  fish  of  the  list  then  the  latter  may  become  infected.  Thus 
it  is  possible  to  offer  explanations  that  will  cover  the  whole  question  of 
the  source  of  infection  for  these  hosts  which  have  such  different  food 
habits.  The  analysis,  however,  is  not  very  convincing.  The  more  plau- 
sible explanation  is  that  several  species  of  parasite  are  being  dealt  with 
under  this  one  name.  It  seems  not  improbable  that  the  parasites  of 
the  cyprinids  may  belong  to  at  least  two  or  probably  three  species  while 
the  jion-cyprinids  may  or  may  not  harbor  any  of  these.  Further  inves- 
tigations on  the  parasites  of  the  hosts  of  this  list  should  be  undertaken 
with  the  view  of  settling  some  of  these  questions. 

CONCLUSIONS 

A  careful  analysis  of  the  extensive  data  presented  in  this  section 
shows  that  species  of  Proteocephalus  may  occur  in  multiple  hosts  in  the 
following  combinations: 

1)  A  species  of  Proteocephalus  may  occur  in  different  host  species 
of  the  same  genus.  Five  species  are  limited  exclusively  to  various  species 
within  the  same  host  genus. 

,2)  A  species  may  occur  in  the  different  genera  of  the  same  family. 
One  case. 

3)  A  species  may  occur  in  the  members  of  closely  allied  genera, 
i.  e.  of  the  same  order.    Four  cases  are  known. 

4)  A  species  may  occur  in  families  of  very  wide  relationships,  i.  e. 
of  different  orders.    There  are  two  cases  of  which  one  is  doubtful. 

It  is  further  shown  in  this  section  that  when  a  species  of  Proteo- 
cephalus occurs  in  multiple  hosts  under  any  of  the  combinations  above 
suggested  the  host  species  have  a  continuous  distribution.  The  only 
apparent  exception  to  this  rule  is  in  the  case  of  AnguUla  vulgaris  of 
Europe  and  A.  chrysypa  of  North  America,  both  harboring  P.  macro- 
cephalus.  In  this  case,  however,  the  exception  is  apparent  and  not  real 
for  these  two  forms  are  perhaps  no  more  than  varieties  of  a  cosmopoli- 
tan species.  Analysis  of  the  data  presented  in  this  section  also  shows 
that  in  many  cases  of  multiple  hosts  the  food  of  the  different  host  spe- 


303]  PROTEOCEPHALIDAE—LA  RUE  303 

cies  is  alike  in  some  or  all  of  its  elements  tho  the  proportions  of  these 
elements  may  differ  widely.  Since  the  infection  of  the  host  is  only 
accomplished  through  the  food  eaten  a  general  statement  may  be  made 
thus:  The  'parasitic  infestation  of  the  host  is  determined  hy  the  food 
eaten.  The  character  of  the  food  is  determined  by  the  environment 
and  largely  by  the  habitat  preferred  by  the  host.  The  apparent  excep- 
tions to  the  above  conclusion  are  found  in  those  cases  where  a  group  of 
species  acting  as  hosts  to  a  single  species  of  parasite  feed  on  very 
different  elements  of  food.  In  each  of  these  cases  the  probability  has 
been  pointed  out  that  there  has  been  a  confusion  of  species  under  one 
name.  If  such  a  confusion  does  not  exist  then  there  is  evidence  of  the 
remarkable  adaptability  of  the  larval  stages  of  the  parasite  to  the  life 
conditions  of  a  large  series  of  invertebrates  and  fish.  Both  of  these 
possibilities  should  be  investigated. 

The  evidence  as  presented  in  this  section  is  not  in  favor  of  a  strict 
adherence  to  the  idea  of  the  specificity  of  parasites.  This  idea  cer- 
tainly does  not  hold  for  any  of  the  better  known  species  of  Proteoceph- 
alus  for  which  quite  a  number  of  collection  records  are  available,  but 
it  seems  to  hold  for  a  number  of  species  for  which  there  are  but  one  or 
two  collection  records  and  for  species  which  are  imperfectly  known. 
To  apply  the  idea  to  these  cases  is  not  justified.  To  break  away  en- 
tirely, however,  from  the  idea  of  specificity  of  parasites  would  be  a 
rash  step,  indeed,  for  certain  cases  have  been  well  established  which 
show  conclusively  that  for  certain  hosts,  other  than  those  considered  in 
this  paper,  certain  species  of  parasites  are  specific.  Such  cases  can 
only  be  determined  by  many  records  of  collection  and  then  are  to  be 
relied  upon  only  after  careful  infection  experiments  have  been  worked 
out.  To  apply  the  idea  of  specificity  of  parasites  without  having  taken 
these  precautions  is  not  justifiable.  In  working  through  the  data  on 
distribution  of  Proteocephalus  species  which  have  multiple  hosts  one 
notes  that  in  many  cases  the  range  of  the  parasite  may  be  greater  than 
the  range  of  some  of  its  hosts.  This  is  well  illustrated  by  the  range  of 
P.  longicollis  which  has  a  far  wider  distribution  than  some  of  the  spe- 
cies of  Coregonus  in  which  it  occurs.  Some  of  these  species  occur  in 
just  a  few  lakes  while  P.  longicollis  is  known  from  many  localities  of 
northern  and  central  Europe.  So  also  the  range  of  P.  macrocephalus 
is  greater  than  that  of  either  of  its  hosts. 

An  explanation  for  this  wide  distribution  of  the  parasite  may  be 
stated  thus:  The  parasite  may  have  a  better  means  of  dispersal  than 
the  host.  The  distribution  of  fish  Proteocephalids  may  be  widened  in 
the  following  ways:  They  are  carried  by  the  primary  host  which 
voiding  feces  as  it  wanders  about  scatters  the  eggs  of  the  parasite  that 


304  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [304 

may  infect  suitable  intermediate  hosts.  The  latter,  vertebrates  or  in- 
vertebrates, may  migrate  or  be  carried  by  currents  to  farther  limits. 
New  hosts  may  be  infected  and  these  on  their  migrations  may  carry  the 
infection  beyond  the  original  limits  of  distribution  of  the  parasite.  The 
introduction  by  man  of  fish  into  new  regions  is  a  method  of  widening 
the  range  of  a  parasite.  It  is  suggested  as  a  further  possibility  that 
fish-eating  birds  at  the  time  of  migration  may  carry  the  eggs  of  the 
parasites  ingested  with  the  fish  for  some  distance  and  these  eggs  being 
discharged  with  the  feces  into  the  water  may  infect  suitable  intermedi- 
ate hosts.  This  method  of  dispersal  could  be  possible  only  if  the  eggs 
of  the  parasite  thus  eaten  are  capable  of  withstanding  the  vigorous 
digestive  action  of  the  bird.  Experiments  would  be  required  to  deter- 
mine the  value  of  this  method. 

Families  op  Fish  from  Which  Proteocephalids  are  Known 
The  data  on  this  subject  are  collected  in  the  accompanying  table. 
It  is  there  shown  that  Proteocephalids  are  known  from  fifteen  families 
of  fish  of  the  group  Teleostomi  and  representing  the  orders  Crossop- 
terygii,  Chondrostei,  Holostei,  and  Teleostei.  The  writer  follows  the 
classification  of  Bridge  (1904)  and  Boulenger  (1904),  Vol.  VII,  Cam- 
bridge Natural  History.  Of  the  fifteen  families  ten  belong  to  the  order 
Teleostei.  The  latter  contains  by  far  the  larger  number  of  fresh  water 
fishes.  Proteocephalid  species  are  as  yet  unknown  from  the  Dipnoi.  A 
comparison  of  the  collection  records  and  of  the  accompanying  table 
shows  that  certain  species  of  parasite  seem  to  be  characteristic  of  cer- 
tain families  of  fish.  Those  ,cases  in  which  the  parasite  has  been  recorded 
but  once  or  twice  probably  should  not  be  considered  as  characteristic 
of  their  host  families.  Among  such  cases  are  to  be  mentioned  the  occur- 
rence of  P.  pentastoma  in  the  Polypteridae,  P.  skorikowi  in  the  Acipen- 
seridae,  P.  singularis  in  Lepisosteidae,  etc.  There  are  certain  species 
which  have  been  recorded  several  times  and  the  majority  of  these 
times  from  a  certain  family  of  fish.  Such  species  are  to  be  considered 
as  being  characteristic  for  the  family.  The  species  which  fall  in  this 
group  are:  P.  longicollis  and  P.  exiguus,  characteristic  of  Salmonidae; 
P.  torulosus  of  the  Cyprinidae ;  P.  macrocephalus  of  Anguillidae ;  P. 
pinguis  of  the  Esocidae;  P.  fUicollis  of  Gasterosteidae ;  P.  ambloplitis 
of  Centrarchidae ;  P.  percae  and  P.  cernuae  of  the  Percidae.  The  re- 
maining species  are  too  little  knoA\Ti  to  justify  an  attempt  to  determine 
this  relationship. 


305] 


PROTEOCEPHALIDAE—LA  RUE 


305 


Families  of  Fish  from  which  Proteocephalids  are  Known,  the  Species  of  the  Latter 
and  the  Frequency  of  their  Occurrence 


Frequence 

Order 

Family 

Species  of  parasite 

of 

occurrence 

Crossopterygii 

Polypteridae 

Proteocephalus  sulcatus 

I 

P.  pentastoma 

2 

Chondrostei 

Acipenseridae 

P.  skorikowi 

I 

Holostei 

Amiidae 

P.  per  plexus 

I 

P.  ambloplitis 

2 

Teleostei 

Lepisosteidae 

P.  singularis 

I 

P.  perplexus 

I 

Clupeidae 

P.  agonis 

I 

Salmon  idae 

P.  longicolHs                            20 

or  more 

P.  fallax 

I 

P.  neglectus 

2(?) 

P.  pusillus 

2 

P.  exiguus 

3 

P.  percae 

I 

P.  salmonis-umblae  (sp.  inq.) 

2 

P.  Cyclops  (Immature  and 

poorly  known) 

I 

P..  salvelini  (Immature  and 

poorly  known) 

I 

P.  torulosus  (Doubtful  deter- 

mination) 

4 

Cyprinidae 

P.  torulosus                                18 

or  more 

P.  sagittus   (sp.  inq.) 

I 

P.  longicollis.     (Doubtful  de- 

termination 

3 

Siluridae 

P.  sulcatus 

I 

P.  fossatus 

I 

P.  osculatus   (sp.  inq.) 

2 

Corallobothrium  solidum 

I 

C.  lobosum 

I 

Choanoscolex  abscisa 

I 

Anguillidae 

Proteocephalus  macrocephalus 

10 

P.  hemisphaericus  (sp.  inq.) 

I 

Esocidae 

P.  esocis 

I 

P.  pinguis 

3 

P.  nematosoma  (sp.  inq.) 

2 

P.  longicollis     (Doubtful  de- 

termination) 

I 

Gastrosteidae 

Proteocephalus  filicollis 

10 

306 


ILLINOIS  BIOLOGICAL  MONOGRAPHS 


[306 


Frequence 

Order 

Family 

Species  of  parasite 

of 

occurrence 

Gadidae 

P.  torulosus  (Doubtful  de- 
termination) 

I 

Centrarchidae 

P.  ambloplitis 

4 

Percidae 

P.  percae 
P.  cernuae 
P.  dubius 

P.  longicollis  (Doubtful  de- 
termination) 

5 
3 

I 

I 

Cottidae 

P.  percae  (needs  confirmation) 

I 

Cichlidae 

P.  macrophallus 

I 

Frequency  of  occurrence  in  the  above  table  has  been  determined  by  taking 
the  sum  of  the  number  of  localities  from  which  the  species  has  been  reported 
(including  times  from  same  locality)  or  in  case  it  occurs  in  more  than  one  host 
in  one  locality  the  number  of  host  species  is  added.     Numbers  are  approximate. 


LIFE   HISTORY  OF  THE  PROTEOCEPHALIDAE 

Since  the  appearance  of  a  previous  paper  by  the  writer  (La  Rue 
1909)  in  which  he  reviewed  the  literature  on  thus  subject  and  in  which 
he  outlined  certain  feeding  experiments  which  had  for  their  object  the 
solution  of  the  problem  of  the  life  history  of  the  cestode  infesting 
Amblystoma  tigrinum  but  little  additional  data  have  appeared  on  the 
life  history  of  Proteocephalids.  Barbieri  (1909)  found  that  Bythro- 
trephes  and  Leptodora  act  as  intermediate  hosts  for  Proteocephalus 
agonis.  So  far  as  can  be  ascertained  there  are  no  other  data  in  the 
recent  literature  of  this  subject.  Fuhrmann  (1903)  outlined  the  life 
history  of  Proteocephalids,  Several  of  the  older  investigators  have 
mentioned  the  finding  of  plerocercoids  in  the  livers  of  fish  which  har- 
bored species  of  Proteocephalus  ia  the  intestine.  The  writer  (La  Rue 
1909)  recorded  the  finding  of  many  plerocercoids  in  the  tissues  of  the 
salamander,  Amblystoma  tigrinum,  and  the  positive  results  obtained 
from  feeding  these  plerocercoids  to  uninfected  hosts.  The  probability 
was  there  suggested  that  salamanders  might  become  infected  through 
the  eating  of  infected  salamanders.  There  is  at  present  no  reason  for 
changing  this  view.  While  making  an  examination  of  a  specimen  of 
Natrix  rhomhifer  the  wTiter  found  numerous  encysted  plerocercoids  in 
the  tissues  of  the  alimentary  tract  and  of  the  liver.    Since  many  snakes 


307]  PROTEOCEPHALIDAE—LA  RUE  307 

are  accidental  or  intentional  cannibals  it  is  possible  that  the  eating  of 
one  snake  by  another  may  be  one  method  of  infection.  Since  also  this 
species  of  snake  lives  largely  upon  fish  and  frogs  one  should  look  to 
these  food  animals  for  the  intermediate  host  of  its  cestodes.  No  avail- 
able records  deal  with  this  method  of  infection. 

From  the  data  presented  by  various  workers  it  seems  probable  that 
the  life  history  of  the  Proteocephalids  is  essentially  as  follows :  The 
eggs  and  some  of  the  ripe  proglottids  bearing  eggs  are  voided  by  the 
host  into  the  water  where  they  are  eaten  by  an  invertebrate,  perhaps  a 
worm,  an  insect  larva,  or  a  crustacean,  or  possibly  the  eater  is  a  verte- 
brate, fish,  snake  or  an  amphibian  of  the  same  species  as  the  host  or 
different.  If  the  invertebrate  or  vertebrate  furnishes  a  suitable  habitat 
for  the  development  of  the  parasite  the  six-hooked  embryo  establishes 
itself  and  from  it  develops  a  plerocercoid  about  which  the  host  produces 
a  cyst.  If  the  intermediate  host  be  eaten  by  a  vertebrate  which  furnishes 
a  proper  habitat  for  the  adult  parasite  the  plerocercoid  when  it  is 
released  by  the  action  of  the  digestive  juices  from  its  intermediate  host 
and  from  its  cyst  passes  to  the  intestine  and  develops  into  the  adult 
tapeworm.  If  the  final  host  engulfs  material  containing  eggs  of  the 
cestode  harbored  by  itself  or  its  congeners  or  perhaps  by  members  of 
other  species  the  host  becomes  infected  with  the  plerocercoids  and  so  it 
may  function  as  a  secondary  as  well  as  a  primary  host  for  its  parasitic 
species.  Cannibalism  may  be  a  means  in  the  spread  of  the  parasites 
harbored.  The  problems  connected  with  the  life-history  of  these  para- 
sites must  ultimately  be  settled  by  experimental  methods. 

ORIGIN  OF  THE  PROTEOCEPHALIDAE 

This  work  has  thrown  some  light  on  the  relations  of  the  genera  and 
species  of  Proteocephalidae  to  one  another.  It  is  of  interest  to  attempt 
to  point  out  the  probable  relations  of  the  family  to  other  cestodes.  Like 
many  other  cestodes  the  members  of  this  family  were  once  included 
in  the  great  genus  Taenia.  That,  however,  was  before  the  internal 
structures  of  these  cestodes  were  well  known  and  it  seems  that  the  basis 
for  this  classification  was  the  apparent  similarity  of  the  suckers.  It  is 
now  evident  that  the  Proteocephalids  are  most  closely  related  struc- 
turally to  the  cestodes  of  the  order  Tetraphyllidea.  A  marked  agree- 
ment in  the  general  arrangement  of  all  the  internal  organs  of  Tetraphyl- 
lidean  cestodes  is  to  be  noted.  In  these  respects  the  Proteocephalids 
do  not  agree  with  the  Cyclophyllidea.  In  external  features  alone  the 
Proteocephalids  resemble  the  Cyclophyllidea  more  than  the  Tetraphyl- 
lidea. This  is  due  to  the  marked  simplicity  of  the  heads  and  suckers  of 
the  Proteocephalids  and  the  usual  hemispherical  form  of  the  suckers. 


N^ 


308  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [308 

The  resemblance  of  the  heads  and  suckers  of  Proteocephalids  to 
those  of  Cyclophyllideans  is,  however,  more  apparent  than  real.  The 
heads  of  the  latter  group  are  not  readily  mobile.  Except  in  those  eases 
where  a  protrusible  rostellum  is  present  the  heads  do  not  change  shape 
much.  The  suckers,  moreover,  are  strongly  cupped  at  all  times  and 
are  not  much  moved  by  the  muscles  of  the  head.  The  heads  of  the 
Proteocephadids  are,  as  the  name  implies,  ever  changing.  They  undergo 
a  multitude  of  changes  of  form  and  appearance.  The  suckers  may  be 
strongly  cupped  when  contracted  or  the  cavity  may  be  entirely  obliter- 
ated by  the  stretching  of  the  sucker.  The  suckers  are  very  mobile, 
being  capable  of  being  pushed  out,  forward  or  backward  or  laterally, 
and  retracted.  This  movement  of  the  suckers  adds  greatly  to  the  protean 
character  of  the  head.  The  whole  anterior  part  of  the  head  may  in  some 
species  be  retracted  within  the  swollen  bulbous  basal  part  of  the  head. 
In  respect  to  the  movement  of  the  head  and  the  action  of  the  suckers 
the  Proteocephalids  resemble  the  other  Tetraphyllideans. 

Another  point  of  resemblance  between  the  Proteocephalids  and  the 
other  Tetraphyllideans  is  the  presence  of  a  degenerate  fifth  sucker  in 
many  species  of  the  two  groups.  In  the  Cyclophyllidea  a  somewhat 
similar  appearing  structure  is  to  be  found  in  certain  species.  Here  it 
is  probably  a  degenerate  rostellum  and  not  a  degenerate  fifth  sucker. 
Structurally  the  Proteocephalidae  are  to  be  considered  as  being  closely 
allied  to  the  Tetraphyllidea  while  their  relationship  to  the  Cyclophyl- 
lidea is  distant.  The  relation  of  the  Proteocephalidae  to  the  other 
groups  of  cestodes  is  remote.  The  members  of  the  order  Tetraphyllidea 
other  than  the  Proteocephalidae  and  Monticellidae  are  characteristic 
i/  parasites  of  marine  fishes, 'mostly  selachians.  The  freshwater  fishes  have 
arisen  from  the  marine  fishes  and  their  parasites  have  doubtless  arisen 
from  the  parasites  of  the  marine  fishes,  the  Proteocephalids  coming  from 
a  Tetraphyllidean  ancestry. 

Just  which  fish  species  was  responsible  for  the  bringing  of  this 
Proteocephalid  stock  into  fresh  water  is  a  matter  of  conjecture.  Since 
this  group  of  parasites  is  found  to  be  parasitic  in  members  of  so  many 
families  of  freshwater  fish  it  may  be  inferred  that  some  of  the  older 
and  more  primitive  Teleostomi,  a  Ganoid,  brought  into  the  freshwater 
environment  a  Tetraphyllidean  with  a  simple  type  of  head  and  suckers, 
altho  this  simplicity  may  have  been  a  later  evolution.  Another  hypothe- 
sis is  that  these  parasites  were  introduced  into  fresh  water  by  representa- 
tives of  several  families  of  fish  such  as  the  Salmonidae,  the  Gadidae,  or 
the  Anguillidae.  Among  the  Salmonidae  certain  species  have  acquired 
the  permanent  habit  of  remaining  in  fresh  water  while  others  enter 
fresh  water  only  to  breed.    The  species  of  AnguiUa  go  back  to  salt  water 


309]  PROTEOCEPHALIDAE—LA  RUE  309 

to  breed  and  hence  each  individual  spends  part  of  its  life  in  a  marine 
environment.  The  Gadidae  are  characteristic  salt  water  fish  altho  the 
members  of  one  genus,  Lota,  have  permanently  established  themselves 
in  the  freshwater  environment.  At  the  present  time  there  is  no  evi- 
dence to  show  that  species  of  Salmonidae  or  of  Anguilla  at  the  time 
of  their  migrations  introduce  into  freshwater  parasitic  species  which  are 
capable  of  establishing  and  maintaining  themselves  in  the  new  environ- 
ment. Ward  (1910)  has  pointed  out  that  Salmo  sehago,  landlocked 
in  lakes  of  eastern  North  America,  has  a  parasitic  fauna  that  is  charac- 
teristic of  its  freshwater  environment.  It  is  evident  then  that  what- 
ever parasites  this  species  brought  with  it  from  its  former  marine 
environment  were  unable  to  establisli  themselves.  This  failure  may 
have  been  due  to  a  lack  of  proper  intermediate  hosts. 

These  considerations  lead  to  a  possible  explanation  which  depends 
on  the  possibility  that  the  intermediate  host  species  (a  fish  or  perhaps 
an  invertebrate)  of  a  Tetraphyllidean  acquired  the  habit  of  living  in 
brackish  and  finally  in  fresh  water.  The  fish  species  which  normally 
made  this  intermediate  host  its  food  may  also  have  made  this  change 
of  environment  or  perhaps  other  fish  began  to  prey  on  the  intermediate 
host.  Under  these  conditions  the  parasite  might  have  become  established 
in  the  new  environment.  It  also  seems  probable  that  new  intermediate 
hosts  might  have  been  acquired.  The  acquiring  of  a  new  host  depends 
not  on  the  anatomical  structures  of  the  host  but  upon  its  physiological 
properties  and  upon  its  habits  of  feeding  and  its  environment.  The 
fact  that  the  head  and  the  suckers  of  Proteocephalus  singularis  strongly 
resemble  these  structures  in  certain  species  of  other  families  of  Tet- 
raphyllidea  is  suggestive.  The  host,  Lepisosteus  platostomus,  is  more- 
over a  primitive  fish  of  strong  carnivorous  habits.  The  Lepisosteidae 
have  long  been  established  in  fresh  water.  It  is  here  suggested  that 
some  member  of  this  family  may  have  been  responsible  for  the  intro- 
duction of  Proteocephalids  into  the  fresh  water  environment.  The  Pro- 
teocephalids  of  amphibians  and  aquatic  reptiles  may  have  been  acquired 
in  the  course  of  development  from  some  fish-like  ancestor  or  it  is  more 
probable  that  they  have  acquired  their  Proteocephalid  fauna  from  eating 
intermediate  hosts  (invertebrates  or  fish)  infected  with  larval  Proteo- 
cephalids. In  this  connection  it  is  of  interest  to  note  that  the  head  of 
Crepidobothrium,  found  in  the  Boidae,  is  the  most  primitive  of  all  the 
heads  of  Proteocephalids  of  amphibians  and  reptiles.  That  is  to  say, 
it  is  more  like  the  heads  of  other  Tetraphyllideans  than  are  the  heads 
of  its  congeners.  The  Boidae  are  considered  to  be  somewhat  primitive 
snakes. 


310  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [310 


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Programma  della  Civica  Scuola  R.  Superiore,  Trieste,  162  pp. 
ViBORG,  E,  N. 

1795.    Sammlung  von   Abhandlungen   fiir  Thierarzte  un   Oekonomen.     Aus 
dem  Danischen.     i.  Bandchen.     324  -j-  2  pp.     Copenhagen. 
VoiGT,  M. 

1903.    Beitrage  zur  Kenntnis   des  Vorkommens  von   Fischparasiten  in   den 
Ploner  Gewassern.     Forschungsber.  biol.  Stat,  zu  Plon,  10 :94-99. 
Wagener,  G.  R. 

1854.     Die    Entwicklung   der   Cestoden.     Nova   acta    Acad.    nat.    curios,   24, 
Suppl.   :2i-9i,  pi.  1-12. 
Ward,  H.  B. 

1894.    A  preliminary  report  on  the  worms    (mostly  parasitic)    collected  in 
Lake  St.  Clair,  in  the  summer  of  1893.    Mich.  Fish  Comm.,  Bull.  4:49-54. 
1910.    Internal  parasites  of  the  Sebago  salmon.     Bull.  U.  S.  Bur.  Fisheries, 
(1908)  28:1151-1194,  pi.  121,  10  fig. 
Wedl,  K. 

1861.    Zur    Helminthenfauna    Aegyptens.     (2.  Abt.)    Sitzungsber.    k.    Akad. 
Wissensch.,  math. -naturw.  CI.,  Wien,  44:463-482,  3  pi.,  42  fig. 
Weinland,  D.  F. 

1858.    Human  cestoides.    x-|-e3  pp.,  12  fig.     Cambridge,  Mass. 
Wright,  R.  R. 

1879.     Contributions    to    American    Helminthology.      No.    i.      Proc.    Canad. 
Institute,  Toronto,  n.  s.,  i  :54-75,  2  pi.,  23  fig.     (Reprint  repaged  3-23.) 
Zeder,  J.  G.  H. 

1800.    Erster    Nachtrag    zur    Naturgeschichte    der    Eingeweidewiirmer,    mit 
Zufassen  und  Anmerkungen  herausgegeben.     xx  +  320  pp.,  6  pi.     Leipzig. 
1803.    Anleitung  zur  Naturgeschichte  der  Eingeweidewiirmer.    xvi  +  432  pp., 
4  pi.     Bamberg. 


318  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [318 

ZSCHOKKE,   F. 

1884.    Recherches  sur  I'organisation   et   la  distribution   zoologique  des  vers 

parasites  des  poissons  d'eau  douce.    Arch.  Biol.,  5:153-241,  pi.  9-10,  16  fig. 

(reprint  repaged  1-89.) 
1888.  '  Recherches  sur  la  structure  anatomique  et  histologique  des  cestodes. 

396  pp.  9  pi.    Geneve. 
1891.     Die    Parasitenfauna   von    Trutta   salar.     Centrbl.    Bakt.    u.    Parasit., 

10 :694-699,  i  pi. ;  738-745 ;  792-801 ;  829-838. 
1896.     Zur    Faunistik    der    parasitischen     Wiirmer    von     Siisswasserfischen. 

Centrbl.  Bakt.  u.  Parasit.,  (i)  19:772-784;  815-825. 
1903.     Marine    Schmarotzer    in     Siissw^asserfischen.     Verhandl.   naturf.    Ge- 

sellsch.  in  Basel,  16:118-157,  pi.  i,  5  fig. 


319] 


PROTEOCEPHALIDAE—LA  RUE 


319 


EXPLANATION  OF  PLATES 

All  original  drawings,  unless  otherwise  stated,  were  drawn  with  the 
aid  of  the  camera  lucida  or  the  Edinger  drawing  apparatus  and  details 
have  been  filled  in  at  the  same  magnification.  Drawings  are  uniformly 
arranged  with  the  anterior  end  or  the  dorsal  surface  toward  the  upper 
end  of  the  plate  unless  there  is  a  statement  to  the  contrary. 


ABBREVIATIONS  USED 


bm 

Basement  membrane 

ci 

Cirrus 

cip 

Cirrus-pouch 

dj 

Ductus  ejaculatorius 

def 

Vas  deferens 

ef 

Vasa  efferentia 

ep 

Epithelium 

exa 

Excretory  anastomosis 

exd 

Excretory  vessel,  dorsal 

exv 

Excretory  vessel,  ventral 

exp 

Excretory  pore 

glc 

Gland  cells 

mc 

Circular  muscles 

mf 

Muscle  fibers 

ml 

Longitudinal  muscles 

mr 

Muscle  rhomboid 

ms 

Muscle  star 

mtr 

Transverse  muscles 

mx 

Muscle  cross 

nl 

Lateral  nerve 

nr 

Nerve  ring 

od  Oviduct 

oot  Ootype 

ov  Ovary 

ooc  Oocapt 

rs  Receptaculum  seminis 

sh  Shell  gland 

tt  Testes 

su  Suckers 

sue  Sucker  cavity 

suf  Sucker,  fifth 

lit  Uterus 

utl  Lateral  uterine  pouches 

utv  Ventral  uterine  outpocketings 

titp  Uterine  passage 

utvp  Ventral  uterine  pores 

va  Vagina 

vas  Sphincter  vaginae 

vi  Vitellaria 

vid  Vitelline  ducts 

vide  Vitelline  duct,  common 


320  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [320 


EXPLANATION   OF   PLATE 

Fig.  ij—Proteocephalus  macrocephalus  (CrepHn),  head,  toto.  Lot  71,  Sebago 
Lake.    Ward  Collection. 

Fig.  2. — Proteocephalus  pusillus  Ward,  head,  toto,  showing  fifth  sucker.  From 
•Cristivomer  namaycush,  Lake  Temagami. 

Fig.  3. — Proteocephalus  pusillus  Ward,  head,  toto,  showing  fifth  sucker.  From 
Salmo  sebago,  Sebago  Lake. 

Fig.  4. — Proteocephalus  pusillus  Ward,  head,  toto,  showing  fifth  sucker.  From 
'Cristivomer  namaycush,  Lake  Temagami. 

Fig.  5. — Proteocephalus  cernuae  (Gmelin),  head,  toto.  From  lot  09.26  Ward 
Collection. 

Fig.  6. — Proteocephalus  cernuae  (Gmelin),  head,  toto.  From  lot  09.26  Ward 
Collection. 

Fig.  7. — Proteocephalus  torulosus   (Batsch),  head,  toto,  a  characteristic  shape. 

Fig.  8. — Proteocephalus  percae  (Miiller),  head,  toto.  From  lot  10.122  Ward 
Collection. 

Fig.  9. — Proteocephalus  percae  (Muller),  head,  toto.  From  lot  10.123  Ward 
Collection. 

Fig.  ID. — Ophiotaenia  perspicua  La  Rue,  head,  toto. 

Fig.  II. — Ophiotaenia  calmettei  (Barrois),  head,  toto. 

Fig.  12. — Crepidobothrium  gerrardii  (Baird),  head,  toto.  From  lot  08.472 
Ward  Collection. 

Fig.  13. — Crepidobothrium  gerrardii  (Baird),  head,  toto.  From  lot  10.190 
Ward  Collection. 


PLATE   I 


321]  PROTEOCEPHALIDAE—LA  RUE  321 


PLATE  II 


322  ILLINOIS  BIOLOGICAL  MONOGRAPHS  U22 


EXPLANATION   OF  PLATE 

Fig.  14. — Proteocephalus  exiguus  La  Rue,  head,  toto. 

Fig.  15. — Proteocephalus  filicollis  (Rud).=:  ambiguus  (Dujardin),  head  drawn 
from  Schneider's  in  toto  preparation. 

Fig.  16. — Proteocephalus  esocis  (Schneider),  head,  drawn  from  a  specimen 
cleared  in  glycerine. 

Fig.  17. — Proteocephalus  perplexus  La  Rue,  head,  toto. 

Fig.  18. — Proteocephalus  amhloplitis  (Leidy),  head,  toto.  From  material  col- 
lected at  Walnut  Lake,  Mich. 

Fig.  19. — Proteocephalus  ambloplitis  (Leidy),  head,  toto.  From  material  col- 
lected at  Walnut  Lake,  Mich. 

Figs.  20,  21  and  22. — Proteocephalus  dubius  La  Rue,  heads,  toto,  showing  fifth 
sucker  and  contraction  states  of  the  suckers. 

Fig.  23. — Proteocephalus  fallax  La  Rue,  head,  toto,  showing  fifth  sucker. 

Fig.  24. — Proteocephalus  singularis  La  Rue,  head,  toto.  Apical  prominence 
contracted. 

Fig.  25. — Proteocephalus  singularis  La  Rue,  head,  toto.  Apical  prominence 
extended. 

Fig.  26. — Ophiotaenia  filaroides  La  Rue,  head  of  adult,  toto.  After  La  Rue  1909. 

Fig.  27. — Ophiotaenia  filaroides  La  Rue,  head  of  plerocercus,  toto,  showing 
vestigial  fifth  sucker  after  atrophy  is  well  under  way.    After  La  Rue  1909, 

Fig.  28. — Ophiotaenia  filaroides  La  Rue,  head  of  plerocercus,  toto,  showing 
outline  of  hypertrophied  fifth  sucker.    After  La  Rue  1909. 

Fig.  29. — Proteocephalus  pmguis  La  Rue,  head,  toto,  dorsal  or  ventral  view. 

Fig.  30. — Proteocephalus  pinguis  La  Rue,  head,  toto,  dorsal  or  ventral  view. 

Fig.  31. — Proteocephalus  pinguis  La  Rue,  head,  toto,  lateral  view.  Figs.  29,  30, 
and  31  are  from  lot  8,  Sebago  Lake.    Ward  Collection. 

Fig.  32. — Proteocephalus  pinguis  La  Rue,  head  and  regions  of  strobila,  toto. 
From  material  collected  at  Walnut  Lake,  Mich. 

Fig.  33. — Crepidobothrium  gerrardii  (Baird),  head,  toto.    After  Smith  1908. 

Fig.  34. — Crepidobothrium  gerrardii  (Baird),  head,  toto.    After  Smith  1908. 

Figs.  35  and  36. — Proteocephalus  pinguis  La  Rue,  heads,  toto,  showing  varia- 
tion in  contraction  states.    From  material  collected  at  Walnut  Lake,  Mich. 

Fig.  37- — Ophiotaenia  marenzelleri   (Barrois),  head,  toto,  after  Schwarz. 

Fig.  38. — Ophiotaenia  grandis  La  Rue,  head,  toto,  showing  swollen  region  back 
of  head. 


PLATE  II 


323]  PROTEOCEPHALIDAE  —  LA  RUE  323 


PLATE  III 


324  ILLINOIS  BIOLOGICAL  MONOGRAPHS  t324 


V 

EXPLANATION  OF  PLATE 

Fig.  39. — Proteocephalus  singttlaris  La  Rue,  head,  cross-section  about  o.io  mm. 
from  the  tip  of  the  head,  near  the  upper  end  of  suckers.  This  section  shows  the 
beginning  of  the  muscle  cross,  mx,  and  the  heavy  muscles  connecting  the  sucker 
wall  with  the  siu-face  of  the  head.  Figs.  39  and  40  are  drawn  to  the  same  scale 
as  Fig.  41. 

Fig.  40. — Proteocephalus  singularis  La  Rue,  head,  cross-section  about  0.135  mm. 
below  tip  of  head  and  about  middle  of  suckers.  Shows  heavy  muscle  across,  mx, 
with  some  fibers  reaching  out  to  the  surface  of  the  head  between  the  suckers. 

Fig.  41. — Proteocephalus  singularis  La  Rue,  head,  cross-section  near  lower 
edge  of  suckers,  showing  muscle  star,  tus,  with  broadly  flared  ends.  Nuclei  are 
shown  as  the  larger  black  spots.  The  closely  packed  groups  of  short  heavy  lines 
are  tangentially  cut  ends  of  muscle  fibers  which  attach  to  the  suckers. 

Fig.  42. — Crepidobothrium  gerrardii  (Baird),  vestigial  fifth  sucker,  cross-sec- 
tion.   Note  sucker  cavity  and  musculature.    Nuclei  could  not  be  seen  in  this  tissue. 

Fig.  43. — Ophiotaenia  filar oides  La  Rue,  vestigial  fifth  sucker,  from  a  frontal 
section  of  a  plerocercus.  Note  the  sucker  cavity  filled  with  granules,  basement 
membrane,  nuclei  and  muscle  fibers  in  and  around  sucker.  Compare  structure  of 
vestigial  sucker  with  that  of  the  normal  sucker  beside  it 

Fig,  44. — Ophiotaenia  filaroides  La  Rue,  vestigial  fifth  sucker,  frontal  section 
of  head  of  plerocercus.  Compare  structures  with  those  of  Fig.  43.  Scale  should 
read  0.05  mm. 

Fig.  45. — Ophiotaenia  filaroides  La  Rue,  vestigial  fifth  sucker  from  a  cross- 
section  of  head  of  a  plerocercus.  Note  the  sucker  cavity  filled  with  granules,  the 
basement  membrane,  the  muscles,  and  nuclei. 

Fig.  46. — Ophiotaenia  filaroides  La  Rue,  vestigial  fifth  sucker,  cross-section  of 
head  of  plerocercus.  Note  points  mentioned  for  Fig.  45.  Sucker  cavity  only 
partly  filled  with  granules.    Scale  should  read  0.05  mm. 


PLATE  III 


325]  PROTEOCEPHALIDAE—LA  RUE  325 


PLATE  IV 


326  ILLINOIS  BIOLOGICAL   MONOGRAPHS  [326 


EXPLANATION   OF   PLATE 

Fig.  47. — Proteocephalus  macrocephalus  (Creplin),  ripe  proglottid  showing  fe- 
male reproductive  organs.    From  Sebago  Lake  material. 

Fig.  48. — Proteocephalus  macrocephalus  (Creplin),  ripe  proglottid,  cross-sec- 
tion in  region  of  cirrus-pouch.  Partial  reconstruction.  From  Sebago  Lake 
material. 

Fig.  49. — Proteocephalus  macrocephalus  (Creplin),  ripe  proglottid  showing 
male  organs.    From  Sebago  Lake  material. 

Fig.  50. — Proteocephalus  exiguus  La  Rue,  mature  proglottid,  toto. 

Fig.  51. — Proteocephalus  exiguus  La  Rue,  end-proglottid,  ventral  view  of  toto 
preparation.    Note  the  excretory  pore  and  the  shape  of  the  ovary. 

Fig.  52. — Proteocephalus  exiguus  La  Rue,  ripe  proglottid,  toto,  as  seen  from 
dorsal  side,  testes  omitted. 

Fig.  53. — Proteocephalus  pusillus  Ward,  ripe  proglottid,  toto,  showing  uterine 
pouches  and  a  rare  abnormality  in  the  position  of  the  vagina  which  is  here  poste- 
rior to  the  cirrus-pouch.     From  Salmo  sebago. 

Fig.  54. — Proteocephalus  pusillus  Ward,  mature  proglottid,  toto.  From  Salmo 
sebago. 

Fig-  55- — Proteocephalus  pusillus  Ward,  mature  proglottid,  toto,  showing  male 
reproductive  organs  and  the  usual  relation  of  vagina  and  cirrus-pouch.  From 
Cristivomer  namaycush. 

Fig.  56. — Proteocephalus  fallax  La  Rue,  ripe  proglottid,  toto.  Shows  two  ven- 
tral uterine  pores.     This  figure  is  inverted. 

Fig.  57- — Proteocephalus  fallax  La  Rue,  mature  proglottid,  toto,  ventral  view. 
Note  musculature  of  cirrus-pouch. 


PLATE  IV 


327]  PROTEOCEPHALIDAE—LA  RUE  327 


PLATE  V 


328  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [328 


EXPLANATION   OF   PLATE 

Fig.  58. — Proteocephalus  filicoUis  (Rud.)  =ambiguus  (Dujardin),  whole  worm. 
This  figure  and  Figs.  59  and  60  were  drawn  from  Schneider's  toto  preparation. 

Fig.  59- — Proteocephalus  filicoUis  (Rud.) ^ambiguus  (Dujardin),  mature  pro- 
glottid, toto. 

Fig.  60. — Proteocephalus  filicoUis  (Rnd.)  =ambiguus  (Dujardin),  ripe  pro- 
glottid, toto. 

Fig.  61. — Proteocephalus  esocis  (Schneider),  young  proglottid,  toto.  Vagina 
and  cirrus-pouch  could  not  be  traced  to  margin  of  proglottid. 

Fig.  62. — Proteocephalus  esocis  (Schneider),  mature  proglottid,  toto. 

Fig.  63. — Proteocephalus  esocis  (Schneider),  outline  of  anterior  end,  posterior 
end  and  middle  of  strobila. 

Fig.  64- — Proteocephalus  perplexus  La  Rue,  ripe  proglottid,  reconstruction 
showing  female  reproductive  organs,  especially  lateral  pouches  of  uterus. 

Fig.  65. — Proteocephalus  perplexus  La  Rue,  ripe  proglottids,  reconstruction 
showing  male  reproductive  organs  and  the  ovary.  Note  also  the  position  of  the 
sphincter  vaginae.  Figs.  64  and  65  are  drawn  from  the  same  series  of  sections. 
Scale  should  read  0.5  mm. 

Fig.  66. — Proteocephalus  cernuae  (Gmelin),  ripe  proglottid,  toto,  showing  es- 
pecially the  uterine  pouches  and  the  slender  ovarian  lobes. 

Fig.  67. — Proteocephalus  cernuae   (Gmelin),  mature  proglottid,  toto. 
Fig.  68. — Proteocephalus  cernuae   (Gmelin),  old  proglottid,  toto. 


PLATE  V 


329]  PROTEOCEPHALIDAE  —  LA  RUE  329 


PLATE  VI 


330  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [330 


EXPLANATION  OF  PLATE 

Fig.  69. — Proteocephalus  percae  (Miiller),  ripe  proglottid,  toto,  ventral  view. 
From  one  of  Schneider's  slides. 

Fig.  70. — Proteocephalus  percae  (Aliiller),  ripe  proglottid,  toto,  dorsal  view. 
Testes  omitted  to  show  uterine  pouches.    From  one  of  Schneider's  slides. 

Fig.  71. — Proteocephalus  percae  (Miiller),  ripe  proglottid,  partial  reconstruc- 
tion from  ventral  sid^  showing  relation  of  reproductive  organs.  From  lot  10.123 
Ward  Collection. 

Fig.  72. — Proteocephalus  percae  (Miiller),  ripe  proglottid,  toto.  Uterine 
pouches  omitted.     From  lot  10.123  Ward  Collection. 

Fig.  73. — Proteocephalus  percae  (Miiller),  fifth  sucker,  section  showing  struc- 
ture.   From  lot  10.122  Ward  Collection. 

Fig.  74. — Proteocephalus  percae  (Miiller),  ripe  proglottid,  partial  reconstruc- 
tion of  cross-sections  to  show  relationship  of  organs  in  region  of  cirrus-pouch  as 
seen  from  anterior  end.  From  lot  10.122  Ward  Collection.  The  letters  va  at  the 
extreme  right  of  the  figure  should  read  vas. 

Fig.  75. — Proteocephalus  dubius  La  Rue,  ripe  proglottid,  toto  showing  uterine 
pouches  and  shape  of  decadent  ovary.    Scale  is  in  error,  each  unit  equals  o.i  mm. 

Fig.  77. — Proteocephalus  dubius  La  Rue,  mature  proglottid,  toto. 


PLATE  \^I^ 


>. 


331]  PROTEOCEPHALIDAE  —  LA  RUE  331 


PLATE  VII 


332  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [332 


EXPLANATION  OF  PLATE 

Fig.  78. — Proteocephalus  torulosus  (Batsch),  ripe  proglottid,  frontal  section 
showing  small  part  of  uterus  and  ovary,  testes  covering  almost  the  entire  area 
between  vitellaria. 

Fig.  79- — Proteocephalus  torulosus  (Batsch),  ripe  proglottid,  frontal  section 
showing  relatively  large  size  of  ovary  and  the  small  number  of  uterine  pouches. 

Fig.  80. — Proteocephalus  torulosus  (Batsch),  ripe  proglottid,  cross-section  in 
region  of  cirrus-pouch. 

Fig.  81. — Proteocephalus  neglectus  La  Rue,  ripe  proglottids,  toto,  showing  re- 
producti\'e  organs.  The  cirrus-pouch  is  much  shortened,  some  ducts  omitted. 
Ventral  view. 

Fig.  82. — Proteocephalus  neglectus  La  Rue,  cirrus-pouch  and  vagina,  normal 
position  and  shape.  Funnel-like  atrium  possibly  a  little  exaggerated.  Note  the 
muscle  fibers  at  end  of  cirrus-pouch. 

Fig.  83. — Proteocephalus  singularis  La  Rue,  ripe  proglottid,  frontal  section 
showing  uterine  pouches  and  the  cut  parts  of  the  much  coiled  excretory  ducts. 

Fig.  84. — Proteocephalus  singularis  La  Rue,  cirrus-pouch  and  vagina,  recon- 
struction. Note  position  of  vaginal  sphincter  and  of  the  dilated  portion  of  the 
vagina. 

Fig.  85. — Proteocephalus  singularis  La  Rue,  ripe  proglottid,  frontal  section 
showing  arrangement  of  testes,  vitellaria  and  ovary.  Vagina  and  cirrus-pouch  not 
shown. 

Fig.  86. — Proteocephalus  singularis  La  Rue,  head,  frontal  section  showing  ap- 
pearance and  position  of  transverse  and  longitudinal  muscles. 

Fig.  87. — Proteocephalus  singularis  La  Rue,  head,  frontal  section  showing 
attachments  of  longitudinal  muscles  to  the  suckers.    Scale  should  read  0.2  mm. 

Fig.  88. — Proteocephalus  pinguis  La  Rue,  proglottid,  portion  of  cross-section 
showing  relation  of  vagina,  cirrus-pouch  and  excretory  ducts. 

Fig.  89. — Proteocephalus  pinguis  La  Rue,  ventral  uterine  pores  leading  to 
exterior.    Shown  in  frontal  section  of  proglottid  just  below  cuticula. 


PLATE  VII 


^]  PROTEOCEPHALIDAE  —  LA  RUE  '    333 


PLATE  VIII 


334  ILLINOIS  BIOLOGICAL  MONOGRAPHS  t334 


EXPLANATION   OF   PLATE 

Fig.  90. — Proteocephalus  pinguis  La  Rue,  cirrus,  vagina,  \as  deferens  and 
portion  of  vasa  efferentia.    From  a  reconstruction.     Scale  should  read  o.i  mm. 

Fig.  91. — Proteocephalus  pinguis  La  Rue,  complex  of  excretory  ducts  leading 
to  the  dorsal  and  ventral  surfaces  of  the  proglottid. 

Fig.  92. — Proteocephalus  pinguis  La  Rue,  ripe  proglottid,  toto,  ventral  view. 
From  lot  8,  Sebago  Lake  material. 

Fig.  93. — Proteocephalus  pinguis  La  Rue,  head,  longitudinal  section  showing 
structure  of  suckers,  etc. 

Fig.  94. — Monticellia  coryphicephala  (Monticelli),  ripe  proglottid,  cross-section 
showing  relations  of  organs  to  the  muscle-sheath.  From  one  of  Monticelli's  prep- 
arations. 

Fig.  95. — Monticellia  coryphicephala  (Monticelli),  ripe  proglottid,  toto,  show- 
ing peculiar  appearance  of  ovary,  distribution  of  testes  over  entire  field  between 
vitellaria,  and  the  scattered  vitelline  follicles.  From  one  of  Monticelli's  prepara- 
tions.    Cf.  Fig.  186. 

Fig.  96. — Monticellia  coryphicephala  (Monticelli),  ripe  proglottid,  cross-section 
through  region  of  ovary  showing  peculiar  relation  of  organs  to  muscle-sheath. 
Note  that  the  ovary  is  partly  outside  and  partly  inside  this  sheath.  From  one  of 
Monticelli's  preparations. 


PLATE  VIII 


33S]  PROTEOCEPHAUDAE—LA  RUE 


PLATE  IX 


336  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [336 


EXPLANATION  OF  PLATE 

Fig.  97. — Ophiotaenia  grandis  La  Rue,  cirrus-pouch  and  vagina,  cirrus  partly- 
protruded. 

Fig.  98. — Ophiotaenia  grandis  La  Rue,  cirrus-pouch  and  vagina,  cirrus  partly 
protruded. 

Fig.  99. — Ophiotaenia  grandis  La  Rue,  organs  of  interovarial  space  as  seen 
in  an  in  toto  preparation. 

Fig.  100. — Ophiotaenia  grandis  La  Rue,  two  proglottids  showing  indentation  at 
the  genital  pore.     No  genital  atrium  visible. 

Fig.  loi. — Ophiotaenia  grandis  La  Rue,  mature  proglottid,  toto,  ventral  view 
showing  reproductive  organs  and  ventral  uterine  pores. 

Fig.  102. — Ophiotaenia  perspicua  La  Rue,  ripe  proglottid,  toto  showing  uterine 
pouches. 

Fig.  103. — Ophiotaenia  filaroides  La  Rue,  ripe  proglottid,  showing  uterine 
pouches  and  protruded  cirrus. 

Fig.  104. — Ophiotaenia  .filaroides  La  Rue,  organs  of  interovarial  space  from 
a  reconstruction.  After  La  Rue  1909.  Through  an  error  in  lettering  the  lower 
part  of  the  vagina  is  marked  'od'  whereas  the  fertilization  passage  beneath  the 
vagina  should  be  marked  'od'.     'Ys*  should  read  'rs'. 

Fig.  105. — Ophiotaenia  filaroides  La  Rue,  mature  proglottid,  toto,  unflat- 
tened,  showing  relationships  of  organs.    After  La  Rue  1909. 


PLATE  IX 


337]  PROTEOCEPHALIDAE  —  LA  RUE  337 


PLATE  X 


338  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [338 


EXPLANATION  OF  PLATE 

Fig.  io6. — Ophiotaenia  trimeresuri  (Parona),  evaginated  cirrus,  cirrus-pouch 
and  vagina.  Note  coils  of  ductus  ejaculatorius  in  the  swollen  cirrus.  Scale  should 
read  0.3  mm. 

Fig.  107. — Ophiotaenia  trimeresuri  (Parona),  cirrus-pouch  and  vagina,  cirrus 
unprotruded.  Note  coils  of  ductus  ejaculatorius  in  the  cirrus-pouch.  Scale  should 
read  0.3  mm. 

Fig.  108. — Ophiotaenia  trimeresuri  (Parona),  mature  proglottid,  toto.  Uterine 
pouches  just  beginning  to  form. 

Fig.  109. — Ophiotaenia  calmettei  (Barrois),  mature  proglottid,  toto.  Uterine 
pouches  more  advanced  than  in  Fig.  no. 

Fig.  no. — Ophiotaenia  calmettei  (Barrois),  mature  proglottid,  toto.  Begin- 
nings of  uterine  pouches  may  be  seen. 

Fig.  III. — Crepidobothrium  gerrardii  (Baird),  mature  proglottid,  toto.  From 
lot  10.179  Ward  Collection. 

Fig.  112. — Crepidobothrium  gerrardii  (Baird),  ripe  proglottid,  toto.  From  lot 
08472  Ward  Collection. 

Fig.  113. — Crepidobothrium  gerrardii  (Baird),  nearly  mature  proglottid,  toto. 
Note  the  great  number  of  testes.     From  lot  10.190  W^ard  Collection. 

Fig.  114. — Crepidobothrium  gerrardii  (Baird),  mature  proglottid,  toto.  Cirrus 
protruded.    Drawn  from  slide  No.  1858,  Bureau  of  Animal  Industrj\  Washington. 

Fig.  115. — Crepidobothrium  gerrardii  (Baird),  mature  proglottid,  toto. 


PLATE  X 


339] 


PROTEOCBPHALIDAE  —  LA  RUB  339 


PLATE  XI 


340  ILLINOIS  BIOLOGICAL  MONOGRAPHS  t340 


EXPLANATION  OF  PLATE 

Fig.  ii6. — Proteocephalus  atnbloplitis  (Leidy),  head,  frontal  section  showing 
vestigial  fifth  sucker.    After  Benedict. 

Fig.  117. — Proteocephalus  atnbloplitis  (Leidy),  head,  toto.    After  Benedict. 

Fig.  118. — Proteocephalus  exiguus  La  Rue,  ripe  proglottid  showing  arrange- 
ment of  organs.    After  Benedict. 

Fig.  119. — Ophiotaenia  lonnbergii  (Fuhrmann),  head,  toto.    After  Fuhrmann. 

Fig.  120. — Proteocephalus  percae  (Miiller),  head,  frontal  section  showing  fifth 
sucker.    After  Schneider. 

Fig.  121. — Proteocephalus  percae  (Miiller),  head,  toto.  Reproduction  of  Miil- 
ler's  figure  2. 

Fig.  122. — Proteocephalus  percae  (Miiller),  head,  toto.  Reproduction  of  Miil- 
ler's  figure  3. 

Fig.  123. — Crepidobothrium  gerrardii  (Baird),  head,  toto.    After  Monticelli. 

Fig.  124. — Crepidobothrium  gerrardii  (Baird),  head,  toto.    After  Monticelli. 

Fig.  125. — Monticellia  macrocotylea  (Monticelli),  head.     After  Monticelli. 

Fig.  126. — Proteocephalus  pentastoma  (Klaptocz),  anterior  face  of  head.  After 
Klaptocz. 

Fig.  127. — Proteocephaluf  pentastoma  (Klaptocz),  head  and  neck.  After  Klap- 
tocz. 

Fig.  128. — Proteocephalus  pentastoma  (Klaptocz),  portion  of  middle  of  worm. 
After  Klaptocz. 

Fig.  129. — Proteocephalus  pentastoma  (Klaptocz),  posterior  part  of  worm. 
After  Klaptocz. 

Fig.  130. — Proteocephalus  sulcatus  (Klaptocz),  head,  anterior  face.  After 
Klaptocz. 

Fig.  131 — Proteocephalus  sulcatus  (Klaptocz),  head.    After  Kloptocz. 
Fig.  132. — Monticellia  coryphicephala  (Monticelli),  head,  toto.    Note  the  prom- 
inence of  the  suckers.    After  Monticelli. 


PLATE  XI 


341]  PROTEOCEPHALIDAE—LA  RUE  341 


PLATE  XII 


342  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [342 


EXPLANATION   OF  PLATE 

Fig.  133. — Proteocephalus  fossatus  (Riggenbach),  head,  toto.  After  Riggen- 
bach. 

Fig.  134a  &  b. — Proteocephalus  ambloplitis  (Leidy),  head  (a)  and  part  of  stro- 
bila  (b).    After  Leidy. 

Fig.  135. — Proteocephalus  exiguus  La  Rue,  entire  worm.    After  Benedict. 

Fig.  136. — Proteocephalus  exiguus  La  Rue,  entire  worm.  After  Benedict. 
These  two  figures  show  the  variation  in  length  of  neck. 

Fig.  137. — Proteocephalus  exiguus  La  Rue,  head,  frontal  section.  After  Bene- 
dict.    Shows  structure  of  the  functional  fifth  sucker. 

Fig.  138. — Proteocephalus  exiguus  La  Rue,  head,  toto  showing  fifth  sucker. 
After  Benedict. 

Fig.  139. — Ophiotaenia  natter eri  (Parona),  head,  toto.    After  Schwarz. 

Fig.  140. — Ophiotaenia  racemosa  (Rud.),  head,  toto.     After  Schwarz. 

Fig.  141. — Ophiotaenia  trimeresuri  (Parona),  head,  toto.     After  Parona. 

Fig.  142. — Ophiotaenia  trimeresuri  (Parona),  head,  toto.     After  Parona. 

Fig.  143. — Proteocephalus  salvelini  (Linton),  head,  toto.     j\fter  Linton. 

Fig.  144. — Proteocephalus  salvelini  (Linton),  head.    After  Linton. 

Figs.  145,  146. — Proteocephalus  torulosus  (Batsch),  heads,  toto.    After  Batsch. 

Figs.  147,  148. — Proteocephalus  dubius  La  Rue,  heads,  toto.    After  Zschokke. 

Fig.  149. — Proteocephalus  dubius  La  Rue,  proglottid.     After  Zschokke. 

Fig.  iso.-^-Proteocephalus  fallax  La  Rue,  head,  toto  showing  fifth  sucker.  After 
Kramer. 

Fig.  151. — Choanoscolex  abscisus  (Riggenbach),  head,  toto.    After  Riggenbach. 
Fig.    152. — Proteocephalus  skorikowi   (von   Linstow),   head,   toto.     After   von 
Linstow. 


137 


140 


3uf. 


152 


139 


136 


135 


144 


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1^  143 


PLATE  XII 


343]  PROTEOCEPHAUDAE  —  LA  RUE  343 


PLATE  XIII 


344  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [344 


EXPLANATION   OF  PLATE 

Fig.  153. — Ophiotaenia  punka  (Cholodkovski),  head,  toto.  After  Cholod- 
kovski. 

Fig.   154. — Monticellia  malopteruri  (Fritsch),  head,  toto.     After  Fritsch. 

Fig.  155. — Ophiotaenia  cahnettei  (Barrois),  head,  toto.    After  Schwarz. 

Fig.  156. — Ophiotaenia  caliiiettei  (Barrois),  head,  toto.     After  Marotel. 

Fig.  157. — Monticellia  diesingii  (Monticelli),  head,  toto.     After  Monticelli. 

Fig.   158. — Proteocephalus  agonis  (Barbieri),  head,  toto.     After  Barbieri. 

Fig.  159. — Proteocephalus  cyclops  (v.  Linstow),  head,  toto.    After  vx)n  Linstow. 

Fig.  160. — Ophiotaenia  pigmentata  (von  Linstow),  head,  toto.  After  von 
Linstow. 

Fig.  161. — Proteocephalus  osculatus  (Goeze),  head,  toto.  From  Cams  after 
Wagener.     This  appears  to  be  a  reproduction  of  Goeze's  figure. 

Fig.  162. — Proteocephalus  osculatus  (Goeze),  young  individual,  toto.  From 
Carus  after  Wagener. 

Fig.  163. — Proteocephalus  osculatus  (Goeze)  =Taenia  alternatiin  transverse 
lineata,  posterior  proglottids,  toto.    After  Goeze. 

Fig.  164. — Proteocephalus  osculatus  (Goeze) =7"a^»ia  alternatini  transverse 
lineata,  head  and  regions  of  strobila.    After  Goeze. 

Fig.  165. — Proteocephalus  osculatus  ( Goeze)  ^=Taenia  alternatini  transverse 
lineata,  head,  toto.    After  Goeze, 


PLATE  XIII 


345]  PROTEOCEPHALIDAE  —  LA  RUE  345 


PLATE  XIV 


34^  ILLINOIS  BIOLOGICAL  MONOGRAPHS  [346 


EXPLANATION  OF  PLATE 

Fig.  i66. — Proteocephalus  tnalopteruri  (Fritsch),  ripe  proglottid.  Modified 
from  Fritsch's  figure. 

Fig.  167. — Proteocephalus  longicollis  (Zeder),  ripe  proglottid.  Modified  from 
drawing  by  von  Linstow. 

Fig.  168. — Proteocephalus  longicollis  (Zeder),  ripe  proglottid,  cross-section. 
After  von  Linstow.    Note  that  the  testes  lie  in  more  than  one  layer. 

Fig.  169. — Proteocephalus  longicollis  (Zeder),  cirrus-pouch.  After  von  Lin- 
stow. 

Fig.  170. — Proteocephalus  fallax  La  Rue,  outline  of  proglottids  from  anterior 
and  posterior  parts  of  strobila.    After  Kramer. 

Fig.  171. — Proteocephalus  iiiacrocephalus  (Creplin)=:7"a<r«xo  dilatata  Linton, 
folds  of  tissue  along  neck.     Afty  Linton. 

Fig.  172. — Proteocephalus  percae  (Miiller),  piece  of  strobila.  Reproduction  of 
Miiller's  figure  4. 

Fig.  173. — Proteocephalus  percae  (Miiller),  strobila.  Reproduction  of  MuUer's 
figure  I. 


PLATE  XIV 


347]  PROTEOCEPHALIDAE—LA  RUE  VO 


PLATE  X\" 


348  ILLINOIS  BIOLOGICAL   MONOGRAPHS  [348 


EXPLANATION   OF  PLATE 

Fig.   174. — iProteocephalus  agonis  (Barbieri),  ripe  proglottid.     After  Barbieri. 

Fig.  175. — Proteocephalus  sulcatus  (Klaptocz),  ripe  proglottid.  After  Klap- 
tocz. 

Fig.  176. — Proteocephalus  sulcatus  (Klaptocz),  proglottids.     After  Klaptocz. 

Fig.  177. — Proteocephalus  skorikowi  (von  Linstow),  ripe  proglottid,  toto. 
After  von  Linstow. 

Fig.  178. — Proteocephalus  fallax  La  Rue,  mature  proglottid,  toto,  showing  re- 
productive organs.     After  Kramer. 

Fig.  179. — Proteocephalus  fallax  La  Rue,  ripe  proglottid,  showing  uterine 
pouches  and  uterine  pore.     After  Kramer. 

Fig.  180. — Proteocephalus  fossatus  (Riggenbach),  proglottid  with  male  organs. 
After  Riggenbach. 

Fig.  181. — MonticelUa  coryphicephala  (Monticelli),  cirrus-pouch  and  vagina. 
After  Monticelli. 

Fig.  182. — Proteocephalus  osculatus  (Goeze),  mature  proglottid.     After  Luhe. 

Fig.  183. — Proteocephalus  ambloplitis  (Leidy),  ripe  proglottid,  frontal  section 
showing  main  parts  of  male  and  female  reproductive  systems.    After  Benedict. 

Fig.  184. — Proteocephalus  torulosus  (Batsch),  ripe  proglottids  showing  organs. 
After  Kramer. 

Fig.  185. — Proteocephalus  agonis  (Barbieri),  mature  proglottid,  uterus  omitted. 
After  Barbieri. 

Fig.  186. — Monticellia  coryphicephala  (Monticelli),  ripe  proglottid,  toto.  After 
Monticelli, 


165 


PLATE  XV 


349]  PROTEOCEPHALIDAE  —  LA  RUE  349 


PLATE  XVI 


350  ILLINOIS  BIOLOGICAL   MONOGRAPHS  fSSO 


EXPLANATION   OF   PLATE 

Fig.  187. — Ophiotaenia  punica  (Cholodkovski),  proglottid,  toto.  After  Cholod- 
kovski. 

Fig.  188. — Ophiotiieiiia  punica  (Cholodkovski),  young  proglottids,  toto.  After 
Cholodkovski. 

Fig.  189. — Ophiotaenia  lonnbergii  (Fuhrmann),  mature  proglottid,  toto.  After 
Fuhrmann. 

Fig.  190. — Crepidobothrium  gerrardti  (Baird),  cirrus  and  vagina.  After  Mon- 
ticelli. 

Fig.  191. — Ophiotaenia  racemosa  (Rud.),  ripe  proglottid,  toto.    After  Schwarz. 

Fig,  192. — Ophiotaenia  trimeresuri  (Parona),  proglottids,  toto.    After  Parona. 

Fig.  193. — Ophiotaenia  trimeresuri  (Parona),  proglottid,  toto.     After  Parona. 

Fig.  194. — Ophiotaenia  nattereri  (Parona),  egg  showing  hooklets  on  outermost 
membrane.    After  Schwarz. 

Fig-  195- — Ophiotaenia  nattereri  (Parona),  ripe  proglottid.     After  Schwarz. 

Fig.  196. — Crepidobothrium  gerrardii  (Baird),  young  proglottids  showing  be- 
ginnings of  reproductive  organs.    After  Smith. 

Fig.  197. — Ophiotaenia  calmettei  (Barrois),  mature  proglottid,  toto.  After 
Marotel. 

Fig.  198. — Ophiotaenia  calmettei  (Barrois),  nearly  ripe  proglottid,  toto.  After 
Marotel. 

Fig.  199. — Ophiotaenia  marenzelleri  (Barrois),  mature  proglottid,  toto.  After 
Schwarz. 


vas 


PLATE  XVI 


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UNIVERSITY  OF  ILLINOIS-URBANA 

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